Carcharodontosauridae
Encyclopedia
Carcharodontosaurids were a group of carnivorous theropod dinosaur
s. In 1931 Ernst Stromer
named Carcharodontosauridae as a family
, in modern paleontology
this name indicates a clade
within Carnosauria
. Carcharodontosaurids included some of the largest land predators ever known: Giganotosaurus
, Mapusaurus
, Carcharodontosaurus
, and Tyrannotitan
all rivaled or exceeded Tyrannosaurus
in size.
, carcharodontosaurids were the largest predators in the early and middle Cretaceous
throughout Gondwana
, with species also present in North America (Acrocanthosaurus
), and Asia (Shaochilong
). Their ages range from the Barremian
(127-121 million years ago) to the Turonian
(93-89 million years ago). However, past the Turonian, there seem to be no signs of the presence of these animals anywhere on the world. They were replaced by the smaller abelisaurids
in Gondwana and by tyrannosaurids
in North America and Asia. According to Fernando Novas
and colleagues, the disappearance of not only carcharodontosaurids but also spinosaurids and other fauna in both Gondwana and North America seem to indicate that this faunal replacement occurred on a global scale.
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), though others place it in the related family Allosauridae (Currie & Carpenter, 2000; Coria & Currie, 2002).
With the discovery of Mapusaurus in 2006, Rodolfo Coria
and Phil Currie
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 giganotosaurine as well.
defined Carcharodontosauridae as a clade
, consisting of Carcharodontosaurus and all species closer to it than to either Allosaurus
, Sinraptor
, Monolophosaurus
, or Cryolophosaurus
. Therefore, this clade is by definition outside of the clade Allosauridae.
The cladogram
below follows the analysis of Brusatte et al., 2009, although note that the placement of Neovenator
has since been revised to outside of but near Carcharodontosauridae, in Neovenatoridae
.
Cladogram after Ortega et al., 2010
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 analysis of Tetanurae
recovered K. petrolicus as a basal carcharodontosaurid in a trichotomy
with Eocarcharia
and a clade comprising more derived
carcharodontosaurids. Bahariasaurus
has also been proposed as a carcharodontosaurid, but its remains are too scarce to be certain. It appears to be synonymous with the ceratosaur
Deltadromeus
.
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 features (see link below). However, these similarities appear to derive from parallel evolution between these two groups. A larger number of cranial and postcranial characters support their relationship with allosaurids.
Dinosaur
Dinosaurs are a diverse group of animals of the clade and superorder Dinosauria. They were the dominant terrestrial vertebrates for over 160 million years, from the late Triassic period until the end of the Cretaceous , when the Cretaceous–Paleogene extinction event led to the extinction of...
s. In 1931 Ernst Stromer
Ernst Stromer
Ernst Freiherr Stromer von Reichenbach was a German paleontologist.He described the following Cretaceous dinosaurs from Egypt: Aegyptosaurus, Bahariasaurus, Carcharodontosaurus, and the largest known theropod, Spinosaurus aegyptiacus...
named Carcharodontosauridae as a family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
, in modern paleontology
Paleontology
Paleontology "old, ancient", ὄν, ὀντ- "being, creature", and λόγος "speech, thought") is the study of prehistoric life. It includes the study of fossils to determine organisms' evolution and interactions with each other and their environments...
this name indicates a clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
within Carnosauria
Carnosauria
Carnosauria is a group of large predatory dinosaurs that lived during the Jurassic and Cretaceous periods. While it originally contained a wide assortment of giant theropods that were not closely related, the group has since been defined to encompass only the allosaurs and their closest kin...
. Carcharodontosaurids included some of the largest land predators ever known: Giganotosaurus
Giganotosaurus
Giganotosaurus is a genus of carcharodontosaurid dinosaur that lived around 97 million years ago during the early Cenomanian stage of the Late Cretaceous Period. It included some of the largest known terrestrial carnivores, slightly larger than the largest Tyrannosaurus, but smaller than the...
, Mapusaurus
Mapusaurus
Mapusaurus was a giant carnosaurian dinosaur from the early Late Cretaceous of what is now Argentina. It was similar in size to its close relative Giganotosaurus, with the largest known individuals estimated as over in length and weighing approximately...
, Carcharodontosaurus
Carcharodontosaurus
Carcharodontosaurus was a gigantic carnivorous carcharodontosaurid dinosaur that lived around 100 to 93 million years ago, during the late Albian to early Cenomanian stages of the mid-Cretaceous Period...
, and Tyrannotitan
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 Giganotosaurus and Carcharodontosaurus...
all rivaled or exceeded Tyrannosaurus
Tyrannosaurus
Tyrannosaurus meaning "tyrant," and sauros meaning "lizard") is a genus of coelurosaurian theropod dinosaur. The species Tyrannosaurus rex , commonly abbreviated to T. rex, is a fixture in popular culture. It lived throughout what is now western North America, with a much wider range than other...
in size.
Evolution
Along with the spinosauridsSpinosauridae
Spinosauridae is a family of specialised theropod dinosaurs. Members of this family were large, bipedal predators with elongated, crocodile-like skulls, sporting conical teeth with no or only very tiny serrations. The front dentary teeth fanned out, giving the animal a characteristic look...
, carcharodontosaurids were the largest predators in the early and middle Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
throughout Gondwana
Gondwana
In paleogeography, Gondwana , originally Gondwanaland, was the southernmost of two supercontinents that later became parts of the Pangaea supercontinent. It existed from approximately 510 to 180 million years ago . Gondwana is believed to have sutured between ca. 570 and 510 Mya,...
, with species also present in North America (Acrocanthosaurus
Acrocanthosaurus
Acrocanthosaurus is a genus of theropod dinosaur that existed in what is now North America during the Aptian and early Albian stages of the Early Cretaceous. Like most dinosaur genera, Acrocanthosaurus contains only a single species, A. atokensis. Its fossil remains are found mainly in the U.S...
), and Asia (Shaochilong
Shaochilong
Shaochilong is a genus of carcharodontosaurid dinosaur from the mid Cretaceous Ulansuhai Formation of China . The type species, S...
). Their ages range from the Barremian
Barremian
The Barremian is an age in the geologic timescale between 130.0 ± 1.5 Ma and 125.0 ± 1.0 Ma). It is a subdivision of the Early Cretaceous epoch...
(127-121 million years ago) to the Turonian
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 93.5 ± 0.8 Ma and 89.3 ± 1 Ma...
(93-89 million years ago). However, past the Turonian, there seem to be no signs of the presence of these animals anywhere on the world. They were replaced by the smaller abelisaurids
Abelisauridae
Abelisauridae is a family of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous Period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found on the modern continents of Africa and South America, as well as on the Indian...
in Gondwana and by tyrannosaurids
Tyrannosauridae
Tyrannosauridae is a family of coelurosaurian theropod dinosaurs which comprises two subfamilies containing up to six genera, including the eponymous Tyrannosaurus. The exact number of genera is controversial, with some experts recognizing as few as three...
in North America and Asia. According to Fernando Novas
Fernando Novas
Fernando Emilio Novas is an Argentine paleontologist working for the Comparative Anatomy Department of the Bernardino Rivadavia Natural History Argentine Museum in Buenos Aires...
and colleagues, the disappearance of not only carcharodontosaurids but also spinosaurids and other fauna in both Gondwana and North America seem to indicate that this faunal replacement occurred on a global scale.
Taxonomy
The family Carcharodontosauridae was originally named by Ernst StromerErnst Stromer
Ernst Freiherr Stromer von Reichenbach was a German paleontologist.He described the following Cretaceous dinosaurs from Egypt: Aegyptosaurus, Bahariasaurus, Carcharodontosaurus, and the largest known theropod, Spinosaurus aegyptiacus...
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), though others place it in the related family Allosauridae (Currie & Carpenter, 2000; Coria & Currie, 2002).
With the discovery of Mapusaurus in 2006, Rodolfo Coria
Rodolfo Coria
Dr. Rodolfo Coria is an Argentine paleontologist.He is best known for having directed the field study and co-naming of the Argentinosaurus in 1993, and the Giganotosaurus , in 1996 among other landmark South American dinosaurs...
and Phil Currie
Phil Currie
Philip John Currie, AOE 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...
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 giganotosaurine as well.
- Family Carcharodontosauridae
- Genus AcrocanthosaurusAcrocanthosaurusAcrocanthosaurus is a genus of theropod dinosaur that existed in what is now North America during the Aptian and early Albian stages of the Early Cretaceous. Like most dinosaur genera, Acrocanthosaurus contains only a single species, A. atokensis. Its fossil remains are found mainly in the U.S...
- Genus CarcharodontosaurusCarcharodontosaurusCarcharodontosaurus was a gigantic carnivorous carcharodontosaurid dinosaur that lived around 100 to 93 million years ago, during the late Albian to early Cenomanian stages of the mid-Cretaceous Period...
- Genus ConcavenatorConcavenatorConcavenator is a genus of theropod dinosaur that lived approximately 130 million years ago during the early Cretaceous period . The type species is C. corcovatus; Concavenator corcovatus means "hump backed hunter from Cuenca"...
- Genus EocarchariaEocarchariaEocarcharia 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 2000 on an expedition led by University of Chicago paleontologist Paul Sereno. The...
- Genus KelmayisaurusKelmayisaurusKelmayisaurus is an extinct genus of carcharodontosaurid theropod dinosaur from the early Cretaceous. Its name refers to the petroleum-producing city of Karamay in the Xinjiang province of western China near where it was found.-Discovery and species:...
- Genus ShaochilongShaochilongShaochilong is a genus of carcharodontosaurid dinosaur from the mid Cretaceous Ulansuhai Formation of China . The type species, S...
- Genus TyrannotitanTyrannotitanTyrannotitan 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 Giganotosaurus and Carcharodontosaurus...
- Subfamily Giganotosaurinae
- Genus GiganotosaurusGiganotosaurusGiganotosaurus is a genus of carcharodontosaurid dinosaur that lived around 97 million years ago during the early Cenomanian stage of the Late Cretaceous Period. It included some of the largest known terrestrial carnivores, slightly larger than the largest Tyrannosaurus, but smaller than the...
- Genus MapusaurusMapusaurusMapusaurus was a giant carnosaurian dinosaur from the early Late Cretaceous of what is now Argentina. It was similar in size to its close relative Giganotosaurus, with the largest known individuals estimated as over in length and weighing approximately...
- Genus Giganotosaurus
- Genus Acrocanthosaurus
Phylogeny
In 1998 Paul SerenoPaul Sereno
Paul Callistus Sereno is an American paleontologist from the University of Chicago who discovered several new dinosaur species on several continents. He has conducted excavations at sites as varied as Inner Mongolia, Argentina, Morocco, and Niger...
defined Carcharodontosauridae as a clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
, consisting of Carcharodontosaurus and all species closer to it than to either Allosaurus
Allosaurus
Allosaurus is a genus of large theropod dinosaur that lived 155 to 150 million years ago during the late Jurassic period . The name Allosaurus means "different lizard". It is derived from the Greek /allos and /sauros...
, Sinraptor
Sinraptor
Sinraptor is a genus of theropod dinosaur from the Late Jurassic. The name Sinraptor comes from the Latin prefix "Sino", meaning Chinese, and "Raptor" meaning thief. The specific name dongi honours Dong Zhiming...
, Monolophosaurus
Monolophosaurus
Monolophosaurus was a genus of theropod dinosaur from the Middle Jurassic of the Shishugou Formation in what is now Xinjiang, China. It was named for the single crest on top of its skull. The type and only known individual is estimated at...
, or Cryolophosaurus
Cryolophosaurus
Cryolophosaurus was a large theropod dinosaur, with a crest on its head that looked like a Spanish comb. Due to the resemblance of this feature to Elvis Presley's pompadour haircut from the 1950s, this dinosaur was at one point informally known as "Elvisaurus".Cryolophosaurus was excavated from...
. Therefore, this clade is by definition outside of the clade Allosauridae.
The cladogram
Cladogram
A cladogram is a diagram used in cladistics which shows ancestral relations between organisms, to represent the evolutionary tree of life. Although traditionally such cladograms were generated largely on the basis of morphological characters, DNA and RNA sequencing data and computational...
below follows the analysis of Brusatte et al., 2009, although note that the placement of Neovenator
Neovenator
Neovenator is a genus of allosauroid dinosaur. Since its discovery on the Isle of Wight, UK, it has become one of the best-known large carnivorous dinosaurs in Europe. Neovenator was at first considered possibly a new species of Megalosaurus. It measured approximately 7.5 meters in length,...
has since been revised to outside of but near Carcharodontosauridae, in Neovenatoridae
Neovenatoridae
Neovenatoridae is a family of large carnivorous dinosaurs. The group is a branch of the allosauroids, a large group of carnosaurs that also includes the sinraptorids, carcharodontosaurids, and allosaurids...
.
Cladogram after Ortega et al., 2010
Tetanurae
Tetanurae, or "stiff tails", is a clade that includes most theropod dinosaurs, as well as birds. Tetanurans first appear during the early or middle Jurassic Period.-Definition:...
recovered K. petrolicus as a basal carcharodontosaurid in a trichotomy
Polytomy
A polytomy , meaning many temporal based branches, is a section of a phylogeny in which the evolutionary relationships can not be fully resolved to dichotomies. In a phylogenetic tree, a polytomy is represented as a node which has more than two immediate descending branches...
with Eocarcharia
Eocarcharia
Eocarcharia 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 2000 on an expedition led by University of Chicago paleontologist Paul Sereno. The...
and a clade comprising more derived
Derived
In phylogenetics, a derived trait is a trait that is present in an organism, but was absent in the last common ancestor of the group being considered. This may also refer to structures that are not present in an organism, but were present in its ancestors, i.e. traits that have undergone secondary...
carcharodontosaurids. Bahariasaurus
Bahariasaurus
Bahariasaurus is a genus of large theropod dinosaur found in the Bahariya Formation in El-Waha el-Bahariya or Bahariya oasis in Egypt and Kem Kem Beds of North Africa, which date to the late Cretaceous Period, , about 95 million years ago...
has also been proposed as a carcharodontosaurid, but its remains are too scarce to be certain. It appears to be synonymous with the ceratosaur
Ceratosauria
Ceratosaurs are members of a group of theropod dinosaurs defined as all theropods sharing a more recent common ancestry with Ceratosaurus than with birds. There is presently no universally agreed upon listing of species or diagnostic characters of Ceratosauria, though they were less derived...
Deltadromeus
Deltadromeus
Deltadromeus is a genus of large basal ceratosaurian theropod dinosaur from Northern Africa. It had long, unusually slender hind limbs for its size, suggesting that it was a swift runner. The skull is not known. Two fossil specimens of a single species Deltadromeus (meaning "delta runner") is a...
.
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 features (see link below). However, these similarities appear to derive from parallel evolution between these two groups. A larger number of cranial and postcranial characters support their relationship with allosaurids.