''Tyrannotitan'' (; ) is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

of huge

biped Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' a ...

carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...

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 ...

of the

carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...

family from the

Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...

stage of the early

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 th ...

period, discovered in

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 ...

. It is closely related to other giant predators like ''

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 ''Carcha ...

'' and especially ''

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 the ...

'' as well as ''

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 be ...

''.

Discovery and species

''Tyrannotitan chubutensis'' was described by Fernando E. Novas, Silvina de Valais, Pat Vickers-Rich, and Tom Rich in 2005. The fossils were found at La Juanita Farm, northeast of Paso de Indios,

Chubut Province Chubut ( es, Provincia del Chubut, ; cy, Talaith Chubut) is a province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa Cruz Province), the Andes ra ...

, Argentina. They are believed to have been from the Cerro Castaño Member,

Cerro Barcino Formation The Cerro Barcino Formation (also known as the Gorro Frigio Formation) is a geological formation in South America whose strata span the Early Cretaceous to the earliest Late Cretaceous. The top age for the formation has been estimated to be Cenoman ...

(

stage Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * Sta ...

). The

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 several ...

material was designated MPEF-PV 1156 and included partial

dentaries In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...

, teeth, back

vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...

e 3–8 and 11–14,

proximal 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 pro ...

tail vertebrae, ribs and

chevrons Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * '' Eulithis testata'', a moth * Chevron (geology), a fold in rock l ...

, a fragmentary

scapulocoracoid The scapulocoracoid is the unit of the pectoral girdle that contains the coracoid and scapula. The coracoid itself is a beak-shaped bone that is commonly found in most vertebrates with a few exceptions. The scapula is commonly known as the ''shoulde ...

humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...

ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...

, partial ilium, a nearly complete

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 with ...

fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity is ...

, and left

metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...

2. Additional material (designated MPEF-PV 1157) included

jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...

s, a right dentary, teeth,

atlas vertebra In anatomy, the atlas (C1) is the most superior (first) cervical vertebra of the spine and is located in the neck. It is named for Atlas of Greek mythology because, just as Atlas supported the globe, it supports the entire head. The atlas i ...

, neck vertebra (?) 9, back vertebrae (?)7, 10, 13, fused sacral centra (5 total), an assortment of

distal 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 pro ...

caudals, ribs, the right femur, a fragmentary left metatarsal 2, pedal

phalanges The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...

2-1, 2–2, and 3-3.

Description

''Tyrannotitan'' was a large animal, reaching in length and in body mass. It is geologically the oldest known giant carcharodontosaurid along with the more basal ''

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, ' ...

'' from North America (both found in Aptian-age rocks). Unlike other known carcharodontosaurids, this animal lacks

skeletal pneumaticity Skeletal pneumaticity is the presence of air spaces within bones. It is generally produced during development by excavation of bone by pneumatic diverticula (air sacs) from an air-filled space, such as the lungs or nasal cavity. Pneumatization is h ...

extending into the sacral and caudal centra. The

is fused, and much better developed than that of ''

'' ''carolinii'', yet the arm is very small. Most of the shaft of the

scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...

is missing. The

acromion In human anatomy, the acromion (from Greek: ''akros'', "highest", ''ōmos'', "shoulder", plural: acromia) is a bony process on the scapula (shoulder blade). Together with the coracoid process it extends laterally over the shoulder joint. The acro ...

curves about 90 degrees from the shaft axis, making it look vaguely

tyrannosaurid 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 genera ...

-like. Whether the sharp difference between taxa is due to evolution or

sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...

in poorly sampled populations of both species, has not been determined (the latter seems unlikely). A proximal caudal has a very tall neural spine (about twice the height of its centrum, judging by the figure). The base of the orbital

fenestra A fenestra (fenestration; plural fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomical st ...

is a notch of nearly 90 degrees into the body of the

, which contrasts with the rounded base restored for ''Giganotosaurus'' and agrees with ''

'' favorably. The denticles on its teeth are "chisel-like", and are virtually identical to those of other carcharodontosaurids in having a wrinkled enamel surface, heavily serrated mesial and distal carinae, and labiolingually compressed (laterally flattened) crowns. The femur of the paratype specimen is long according to Novas ''et al.'' Canale ''et al.'' recover ''Tyrannotitan'' as deeply nested within the tribe Giganotosaurini as its most basal member. Characteristics that unite the Giganotosaurini include the presence of a postorbital process on the jugal with a wide base, and a derived femur with a weak fourth trochanter and a shallow broad extensor groove at the distal end.

Classification

The following cladogram after Novas ''et al.'', 2013, shows the position of ''Tyrannotitan'' within Carcharodontosauridae.

References

{{Portal bar, Dinosaurs, Paleontology, Cretaceous Carcharodontosaurids Aptian life Early Cretaceous dinosaurs of South America Cretaceous Argentina Fossils of Argentina Cerro Barcino Formation Fossil taxa described in 2005 Taxa named by Fernando Novas Taxa named by Patricia Vickers-Rich Taxa named by Tom Rich