Carcharodontosauridae
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Carcharodontosauridae (carcharodontosaurids; from 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 ...
καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous
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 ...
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 ...
s. In 1931,
Ernst Stromer Ernst Freiherr Stromer von Reichenbach (12 June 1871 in Nürnberg – 18 December 1952 in Erlangen) was a German paleontologist. He is best remembered for his expedition to Egypt, during which the first known remains of ''Spinosaurus'' we ...
named Carcharodontosauridae as a
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
, which, in modern
paleontology 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 ...
, indicates a
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 ter ...
within
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 ...
. Carcharodontosaurids include some of the largest land predators ever known: ''
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 ...
'', ''
Mapusaurus ''Mapusaurus'' () was a giant carcharodontosaurid carnosaurian dinosaur from the early Late Cretaceous (early Turonian stage), approximately 93.9 to 89.6 million years ago, of what is now Argentina. Discovery ''Mapusaurus'' was excavated b ...
'', ''
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 ...
'', and ''
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 ...
'' all rivaled or exceeded ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropoda, 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. ''Tyrannosa ...
'' in size. A 2015 paper by Christophe Hendrickx and colleagues gives a maximum length estimate of for the largest carcharodontosaurids, while the smallest carcharodontosaurids were estimated to have been at least long.


Evolution

Along with the
spinosaurids The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...
, carcharodontosaurids were the largest predators in the early and middle
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 ...
throughout
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final sta ...
, with species also present in North America (''
Acrocanthosaurus ''Acrocanthosaurus'' ( ; ) is a genus of carcharodontosaurid dinosaur 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, ' ...
''), Europe (''
Concavenator ''Concavenator'' is an extinct genus of theropod dinosaur that lived approximately 130 million years ago during the early Cretaceous period (Barremian age). The type species is ''C. corcovatus''; ''Concavenator corcovatus'' means " Cuenca hunter ...
'') and Asia (''
Shaochilong ''Shaochilong'' (meaning "shark toothed dragon") is an extinct genus of carcharodontosaurid dinosaur from the mid-Cretaceous (Late Cenomanian to the end of the Turonian stage) Ulansuhai Formation of China (about 92 million years ago). The type ...
''). Carcharodontosaurids range throughout the Cretaceous from the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is preceded ...
(127-121 million years ago) to the
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 ...
(93-89 million years ago). Past the Turonian, they were replaced by the smaller abelisaurids in Gondwana and by
tyrannosaurids Tyrannosauridae (or tyrannosaurids, meaning " tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to thirteen genera, including the eponymous '' Tyrannosaurus''. The exact number of ...
in North America and Asia.Novas, de Valais, Vickers-Rich, and Rich. (2005). "A large Cretaceous theropod from Patagonia, Argentina, and the evolution of carcharodontosaurids." ''Naturwissenschaften'', While some teeth and a
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. T ...
discovered in
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 inte ...
deposits of
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
. this identification has been subsequently rejected and the material assigned to abelisaurids after better examination, and there are no reliable records of carcharodontosaurs in South America beyond the end of the Turonian. In December 2011, Oliver W. M. Rauhut described a new genus and species of carcharodontosaurid from the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
(late
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxford ...
to earliest
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by ...
faunal stage In chronostratigraphy, a stage is a succession of rock strata laid down in a single age on the geologic timescale, which usually represents millions of years of deposition. A given stage of rock and the corresponding age of time will by convent ...
, about 154-150 
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 ...
) of
Tendaguru Formation The Tendaguru Formation, or Tendaguru Beds are a highly fossiliferous formation and Lagerstätte located in the Lindi Region of southeastern Tanzania. The formation represents the oldest sedimentary unit of the Mandawa Basin, overlying Neoproter ...
, southeastern
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...
. This genus, ''
Veterupristisaurus ''Veterupristisaurus'' is an extinct genus of carcharodontosaurid theropod dinosaur known from the Jurassic of Tendaguru, Lindi Region of southeastern Tanzania. Discovery and naming ''Veterupristisaurus'' is known from the holotype specimen ...
'' represents the oldest known carcharodontosaurid.


Classification

The family Carcharodontosauridae was originally named by
Ernst Stromer Ernst Freiherr Stromer von Reichenbach (12 June 1871 in Nürnberg – 18 December 1952 in Erlangen) was a German paleontologist. He is best remembered for his expedition to Egypt, during which the first known remains of ''Spinosaurus'' we ...
in 1931 to include the single newly discovered species ''Carcharodontosaurus saharicus''. A close relative of ''C. saharicus'', ''Giganotosaurus'', was added to the family when it was described in 1995. Additionally, many paleontologists have included ''Acrocanthosaurus'' in this family (Sereno et al. 1996, Harris 1998, Holtz 2000, Rauhut 2003, Eddy & Clarke, 2011, Rauhut 2011), though others place it in the related family Allosauridae (Currie & Carpenter, 2000; Coria & Currie, 2002). Carcharodontosaurids are characterized by the following morphological characters : Dorsoventral depth of anterior maxillary interdental plates more than twice anteroposterior width, squared, sub-rectangular anterior portion of the dentary, teeth with wrinkled enamel surfaces, presence of four premaxillary alveoli and a premaxillary body taller than long in lateral aspect, opisthocoelous cervical vertebrae with neural spines more than 1.9 times the height of the centrum, large, textured rugosities on the lacrimal and postorbital formed by roofing and forming broad orbital shelves, and a proximomedially inclined femoral head. With the discovery of ''Mapusaurus'' in 2006,
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 ...
erected a subfamily of Carcharodontosauridae, the Giganotosaurinae, to contain the most advanced South American species, which they found to be more closely related to each other than to the African and European forms. Coria and Currie did not formally refer ''Tyrannotitan'' to this subfamily, pending a more detailed description of that genus, but noted that based on characteristics of the femur, it may be a gigantosaurine as well. In 1998
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at si ...
defined Carcharodontosauridae as a
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 ter ...
, consisting of ''Carcharodontosaurus'' and all species closer to it than to either ''
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 ...
'', '' Sinraptor'', '' Monolophosaurus'', or ''
Cryolophosaurus ''Cryolophosaurus'' ( or ; "CRY-oh-loaf-oh-SAWR-us") is a genus of large theropod dinosaur known from only a single species ''Cryolophosaurus ellioti'', from the early Jurassic of Antarctica. It was one of the largest theropods of the Early Jura ...
''. Therefore, this clade is by definition outside of the clade Allosauridae. The
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 ...
below follows the analysis of Brusatte ''et al.'', 2009. Cladogram after Ortega ''et al.'', 2010 Cladogram after Novas ''et al.'', 2013 Cladogram after Canale ''et al.'', 2022 The placement of ''Acrocanthosaurus'' is unclear, with most researchers favoring Carcharodontosauridae and others favoring Allosauridae. In 2011, a redescription of ''Kelmayisaurus'' by Stephen L. Brusatte, Roger B. J. Benson and Xing Xu found it to be valid genus of Carcharodontosauridae. A
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
analysis of
Tetanurae Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, compsognathids and maniraptorans (including birds). Tetanurans ar ...
recovered ''K. petrolicus'' as a basal carcharodontosaurid in a trichotomy with ''
Eocarcharia ''Eocarcharia'' (meaning "dawn shark") is a genus of carcharodontosaurid theropod dinosaur from the Early Cretaceous Elrhaz Formation that lived in the Sahara 112 million years ago, in what today is the country of Niger. It was discovered in 2 ...
'' and a clade comprising more
derived Derive may refer to: *Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguation ...
carcharodontosaurids. ''
Bahariasaurus ''Bahariasaurus'' (meaning " Bahariya lizard") is an enigmatic genus of large theropod dinosaur. ''Bahariasaurus'' is known to have included at least 1 species, ''Bahariasaurus ingens'', which was found in North African rock layers dating to the ...
'' has also been proposed as a carcharodontosaurid, but its remains are too scarce to be certain. Carcharodontosaurids have been proposed as more closely related to abelisaurids, as opposed to the allosaurids. This is due to these two clades sharing some
cranial Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
features. However, these similarities appear to derive from parallel evolution between these two groups. A larger number of cranial and
postcranial Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated s ...
characters support their relationship with allosaurids.


Paleobiology


Growth

Studies of the Argentinian taxon ''
Meraxes ''Meraxes'' is a genus of carcharodontosaurid theropod dinosaur from the Late Cretaceous Huincul Formation of Argentine Patagonia. The genus contains a single species, ''Meraxes gigas''. Discovery and naming The holotype of ''Meraxes'', prev ...
'' suggest that these allosauroids were slow-growing, taking as long as 30-40 years to reach maturity. The maximum age for the holotype specimen was estimated to be roughly 50 years at the time of its death, making it the oldest known non-avian theropod. Unlike ''Tyrannosaurus'', to which its growth has been compared, this carcharodontosaurid was revealed to have kept growing throughout its life. In 2021 the paleontologist Andrea Cau proposed that
Scipionyx ''Scipionyx'' ( ) was a genus of theropod dinosaur from the Early Cretaceous of Italy, around 113 million years ago. There is only one fossil known of ''Scipionyx'', discovered in 1981 by an amateur paleontologist and brought to the attention ...
, formally classified as a
compsognathid Compsognathidae is a family of coelurosaurian theropod dinosaurs. Compsognathids were small carnivores, generally conservative in form, hailing from the Jurassic and Cretaceous Periods. The bird-like features of these species, along with other d ...
, was instead an hatchling carcharodontosaurid. He compared it with
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 ...
and
Acrocanthosaurus ''Acrocanthosaurus'' ( ; ) is a genus of carcharodontosaurid dinosaur 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, ' ...
, finding it more similar to the carcharodontosaurid than the
allosaurid Allosauridae is a family of medium to large bipedal, carnivorous allosauroid theropod dinosaurs from the Late Jurassic. Allosauridae is a fairly old taxonomic group, having been first named by the American paleontologist Othniel Charles Marsh in ...
.


References


External links


Carcharodontosauridae/Abelisauridae
{{Taxonbar, from=Q131334 Carcharodontosauria Kimmeridgian first appearances Turonian extinctions Prehistoric dinosaur families