Hubbry Logo
CarnosauriaCarnosauriaMain
Open search
Carnosauria
Community hub
Carnosauria
logo
8 pages, 0 posts
0 subscribers
Be the first to start a discussion here.
Be the first to start a discussion here.
Carnosauria
Carnosauria
Carnosauria
View on Wikipedia
from Wikipedia

Not to be confused with Carnotaurus.
"Carnosaur" redirects here. For the 1984 horror novel, see Carnosaur (novel). For the 1993 film adaptation, see Carnosaur (film). For the film series, see Carnosaur (series).

Carnosaurs
Temporal range:
Middle Jurassic – Late Cretaceous, 174–90 Ma Possible Toarcian records.
Four carnosaurs (top to bottom): Sinraptor, Acrocanthosaurus, Concavenator, Allosaurus
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Avetheropoda
Clade: †Carnosauria
von Huene, 1920
Subgroups
Synonyms

Allosauroidea? Marsh, 1878

Carnosauria is an extinct group of carnivorous theropod dinosaurs that lived during the Jurassic and Cretaceous periods.

While Carnosauria was historically considered largely synonymous with Allosauroidea, some recent studies have revived Carnosauria as clade including both Allosauroidea and Megalosauroidea (which is sometimes recovered as paraphyletic with respect to Allosauroidea), and thus including the majority of non-coelurosaurian members of theropod clade Tetanurae.[1] Other researchers have found Allosauroidea and Megalosauroidea to be unrelated groups.[2]

Distinctive characteristics of carnosaurs include large eye sockets, a long narrow skull and modifications of the legs and pelvis such as the thigh (femur) being longer than the shin (tibia).[3]

Carnosaurs first appeared in the Middle Jurassic around 174 million years ago, and the last definitive carnosaur family Carcharodontosauridae became extinct in the Turonian epoch of the Late Cretaceous around 90 million years ago. Some theropod remains, once suggested to be putative carcharodontosaurids from the Maastrichtian epoch (72–66 mya) in South America, were later reinterpreted as those of other theropod groups including the abelisaurids and maniraptorans.[4][5]

History of study

[edit]

Carnosauria has traditionally been used as a dumping ground for all large theropods. Even non-dinosaurs, such as the rauisuchian Teratosaurus, were once considered carnosaurs. However, analysis in the 1980s and 1990s revealed that other than size, the group shared very few characteristics, making it polyphyletic. Most former carnosaurs (such as the megalosaurids, the spinosaurids, and the ceratosaurs) were reclassified as more primitive theropods. Others (such as the tyrannosaurids) that were more closely related to birds were placed in Coelurosauria. Modern cladistic analysis defines Carnosauria as those tetanurans sharing a more recent common ancestor with Allosaurus than with modern birds.[6]

Anatomy

[edit]
Size comparison of seven carnosaurs

Carnosaurs share certain distinctive features, one of which is a triangular-shaped pubic boot.[7] They also have 3 fingers per hand, with the second and third digit being approximately equal in length. The femur is larger than the tibia. Another defining feature of carnosaurs is that the chevron bases on their tails have anterior and posterior bone growth.[8] The largest carnosaurs can reach up to 10 meters in length. The length of the body from the tail to the hip is between 54% and 62% of the total body length, and the length of the body from the head to the hip is between 38% and 46% of the total body length.[9] Carnosaurs scaled their limbs relative to their body in a way similar to how other large theropods, like the tyrannosaurids, did.[10] During the Cretaceous, some carnosaurs grew to sizes similar to those of the largest tyrannosaurids.[11] These large carnosaurs lived in the same time period as the other large theropods found in the upper Morrison and Tendaguru formations.[12]

Carnosaurs maintained a similar center of mass across all sizes, which is found to be between 37% and 58% of the femoral length anterior to the hip. Other similarities across all carnosaurs include the structure of their hind limb and pelvis. The pelvis in particular is thought to be designed to reduce stress regardless of body size. In particular, the way the femur is inclined reduces the bending and torsion stress. Furthermore, like other animals with tails, carnosaurs possess a caudofemoralis longus (CFL) muscle that allowed them to flex theirs. Larger carnosaurs are found to have a lower CFL muscle-to-body-mass proportion that smaller carnosaurs.[9]

In addition to body similarities, most carnosaurs, especially most allosauroids are also united by certain skull features. Some of the defining ones include a smaller mandibular fenestra, a short quadrate bone, and a short connection between the braincase and the palate.[13] Allosauroid skulls are about 2.5 to 3 times longer as they are tall.[12] Their narrow skull along with their serrated teeth allow carnosaurs to better slice flesh off of their prey. Carnosaur teeth are flat and have equally-sized denticles on both edges. The flat side of the tooth face the sides of the skull, while the edges align on the same plane as the skull.[14] From analyzing the skull of different carnosaurs, the volume of the cranial vault ranges between 95 milliliters in Sinraptor to 250 milliliters in Giganotosaurus.[15]

Allosaurus and Concavenator preserve skin impressions showing their integument. In Allosaurus, skin impressions showing small scales measuring 1 to 3 millimeters are known from the side of the torso and the mandible. Another skin impression from the ventral side of the neck preserves broad scutate scales. An impression from the base of the tail preserves larger scales around 2 centimeters in diameter. However, it has been noted that these may be sauropod scales due to their similarity and the fact that non-theropod remains were discovered associated with the tail of this particular Allosaurus specimen.[16] Concavenator preserves rectangular scutate scales on the underside of the tail, as well as scutate scales on top of the feet along with small scales on the rest of the feet. A series of knobs on the ulna of Concavenator have been interpreted by some authors as quill knobs theorized to have supported primitive quills;[17] however this interpretation has been questioned, and they have been suggested to represent traces of ligaments instead.[18]

Classification

[edit]

Within Carnosauria, there is a slightly more exclusive clade, superfamily Allosauroidea. The clade Allosauroidea was originally named by Othniel Charles Marsh, but it was given a formal definition by Phil Currie and Zhao, and later used as a stem-based taxon by Paul Sereno in 1997.[19] Sereno was the first to provide a stem-based definition for the Allosauroidea in 1998, defining the clade as "All neotetanurans closer to Allosaurus than to Neornithes."[20] Kevin Padian used a node-based definition in his 2007 study which defined the Allosauroidea as Allosaurus, Sinraptor, their most recent common ancestor, and all of its descendants. Thomas R. Holtz and colleagues and Phil Currie and Ken Carpenter, among others, have followed this node-based definition.[21][22] Depending on the study, Carnosauria and Allosauroidea are sometimes considered synonymous. In such cases, several researchers have elected to use Allosauroidea over Carnosauria.[12][23]

Conventional phylogeny

[edit]

The following cladogram illustrates the phylogenetic position of Allosauroidea within Theropoda. It is a simplified version of the tree presented in a synthesis of the relationships of the major theropod groups based on various studies conducted in the 2010s. The ⊞ button can be clicked to expand the clade and display the interrelationships of the four major allosauroid groups.[24]

Neotheropoda

Coelophysoidea

Dilophosauridae

Averostra

Ceratosauria

Tetanurae

Megalosauroidea

Avetheropoda
Allosauroidea
⊞
⊟
Coelurosauria

Alternative hypotheses

[edit]

The composition of the clade Carnosauria has been controversial among scientists since at least 2010. Different clades have been recovered by different authors, and a scientific consensus has yet to emerge.

One such clade is Neovenatoridae, a proposed clade of carcharodontosaurian carnosaurs uniting some primitive members of the group such as Neovenator with the Megaraptora, a group of theropods with controversial affinities. Other studies recover megaraptorans as basal coelurosaurs unrelated to carcharodontosaurs. Other theropods with uncertain affinities such as Gualicho, Chilantaisaurus and Deltadromeus are also sometimes included.[25][26]

Neovenatoridae, as formulated by these authors, contained Neovenator, Chilantaisaurus, and a newly named clade: Megaraptora. Megaraptora contained Megaraptor, Fukuiraptor, Orkoraptor, Aerosteon, and Australovenator. These genera were allied with the other neovenatorids on the basis of several features spread out throughout the skeleton, particularly the large amount of pneumatization present. The pneumatic ilium of Aerosteon was particularly notable, as Neovenator was the only other taxon known to have that trait at the time. Neovenatorids were envisioned as the latest-surviving allosauroids, which were able to persist well into the Late Cretaceous due to their low profile and coelurosaur-like adaptations.[23] Later studies supported this hypothesis, such as Carrano, Benson & Sampson large study of tetanuran relationships in 2012,[27] and Zanno & Makovicky description of the newly discovered theropod Siats in 2013, which they placed within Megaraptora. Fukuiraptor and Australovenator were consistently found to be close relatives of each other; this was also the case for Aerosteon and Megaraptor. Orkoraptor was a "wildcard" taxon difficult to place with certainty.[28]

Phylogenetic studies conducted by Benson, Carrano and Brusatte (2010) and Carrano, Benson and Sampson (2012) recovered the group Megaraptora and a few other taxa as members of the Neovenatoridae. This would make neovenatorids the latest-surviving allosauroids; at least one megaraptoran, Orkoraptor, lived near the end of the Mesozoic era, dating to the early Maastrichtian stage of the latest Cretaceous period, about 70 million years ago.[23][12]

The cladogram below follows a 2016 analysis by Sebastián Apesteguía, Nathan D. Smith, Rubén Juarez Valieri, and Peter J. Makovicky based on the dataset of Carrano et al. (2012).[29]

Allosauroidea

Subsequent analyses have contradicted the above hypothesis. Novas and colleagues conducted an analysis in 2012 which found that Neovenator was closely related to carcharodontosaurids, simultaneously found Megaraptor and related genera to be coelurosaurs closely related to tyrannosaurids.[30] However, Novas et al. subsequently found that megaraptorans lacked most of the key features in the hands of derived coelurosaurs including Guanlong and Deinonychus. Instead, their hands retain a number of primitive characteristics seen in basal tetanurans such as Allosaurus. Nevertheless, there are still a number of other traits that support megaraptorans as members of the Coelurosauria.[31][32] Other taxa like Deltadromeus and Gualicho have been alternatively recovered as coelurosaurs or noasaurid ceratosaurs.[29] Over the recent years, a majority of researchers have increasingly classified megaraptorans as coelurosaurs.[33][34][35][36]

Several recent analyses do not find a relationship between Neovenator and megaraptorans, which suggests that the latter were not carnosaurs or allosauroids. As a result of these findings, and the fact that Neovenator itself is the only uncontroversial neovenatorid, the family Neovenatoridae sees little use in recent publications.[37][32]

In 2019, Rauhut and Pol described Asfaltovenator vialidadi, a basal allosauroid displaying a mosaic of primitive and derived features seen within Tetanurae. Their phylogenetic analysis found traditional Megalosauroidea to represent a basal grade of carnosaurs, paraphyletic with respect to Allosauroidea. Because the authors amended the definition of Allosauroidea to include all theropods that are closer to Allosaurus fragilis than to either Megalosaurus bucklandii or Neornithes, the Piatnitzkysauridae was found to fall within Allosauroidea. A cladogram displaying the relationships they recovered is shown below.[1]

Carnosauria

In 2025, photos of a destroyed specimen of Carcharodontosaurus were re-examined by Maximilian Kellermann and colleagues and were used to erect the new genus Tameryraptor. A byproduct of this study was the recovery of a novel phylogenetic arrangement of carnosaurs. They did not recover megalosauroids as close relatives of allosauroids. Within Allosauroidea, their analyses, based on the dataset published by Wang and colleagues in 2017, consistently found Metriacanthosauridae and Carcharodontosauridae as sister taxa. They named this novel clade Carcharodontosauriformes and defined it as the least-inclusive clade containing both Carcharodontosaurus and Sinraptor. A simplified version of one of the cladograms they published are shown below.[38]

Allosauroidea

Paleobiology and behavior

[edit]
Allosauroidea skull comparison

Multiple severe injuries have been found on allosauroid remains, which implies that allosauroids were frequently in dangerous situations and supports the hypothesis of an active, predatory lifestyle. Despite the multitude of injuries, only a few of those injuries show signs of infection. For those injuries that did become infected, the infections were usually local to the site of the injury, implying that the allosauroid immune response was able to quickly stop any infection from spreading to the rest of the body. This type of immune response is similar to modern reptilian immune responses; reptiles secrete fibrin near infected areas and localize the infection before it can spread via the bloodstream.[39]

The injuries were also found to be mostly healed. This healing may indicate that allosauroids had an intermediate metabolic rate, similar to non-avian reptiles, which means they require fewer nutrients in order to survive. A lower nutrient requirement means allosauroids do not need to undertake frequent hunts, which lowers their risk of sustaining traumatic injuries.[39]

Although the remains of other large theropods like tyrannosaurids bear evidence of fighting within their species and with other predators, the remains of allosauroids do not bear much evidence of injuries from theropod combat. Most notably, despite a good fossil record, allosauroid skulls lack the distinctive face-biting wounds that are common in tyrannosaurid skulls, leaving open the question of if allosauroids engaged in interspecies and intraspecies fighting.[40] Remains of the allosauroid Mapusaurus are also often found in groups, which could imply the existence of social behavior.[41] While there are alternative explanations for the groupings, like predator traps or habitat reduction due to drought, the frequency of finding allosauroid remains in groups supports the social animal theory. As social animals, allosauroids would share the burden of hunting, allowing injured members of the pack to recover faster.[39]

Paleobiogeography

[edit]

The paleobiogeographical history of allosauroids closely follows the order that Pangaea separated into the modern continents.[42] By the Middle Jurassic period, tetanurans had spread to every continent and diverged into the allosauroids and the coelurosaurs.[11] Allosauroids first appeared in the Middle Jurassic period and were the first giant taxa (weighing more than 2 tons) in theropod history. Along with members of the superfamily Megalosauroidea, the allosauroids were the apex predators that occupied the Middle Jurassic to the early Late Cretaceous periods.[43] Allosauroids have been found in North America, South America, Europe, Africa, and Asia.[42] A probable carcharodontosaurian specimen from the upper Strzelecki Group and the Eumeralla Formation (Aptian-Albian) of Australia might potentially extend their known distribution.[44]

Specifically, a world-wide dispersal of carcharodontosaurids likely happened in the Early Cretaceous. It has been hypothesized that the dispersal involved Italy's Apulia region (the "heel" of the Italian peninsula), which was connected to Africa by a land bridge during the Early Cretaceous period; various dinosaur footprints found in Apulia support this theory.[45] Allosauroids were present in both the northern and southern continents during the Jurassic and Early Cretaceous, but they were later displaced by the tyrannosauroids in North America and Asia during the Late Cretaceous. This is likely due to regional extinction events, which, along with increased species isolation through the severing of land connections between the continents, differentiated many dinosaurs in the Late Cretaceous.[11]

See also

[edit]

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Carnosauria

View on Grokipedia
from Grokipedia
Carnosauria is an extinct of large-bodied carnivorous theropod dinosaurs, phylogenetically defined as Allosaurus fragilis and all taxa sharing a more recent common ancestor with than with Neornithes (modern birds). This group, nested within the larger as part of Avetheropoda, represents some of the most iconic apex predators of ecosystems, sister to the lineage that includes tyrannosaurids and birds. Carnosaurs first appeared in the around 168 million years ago, with the earliest known member being Monolophosaurus jiangi from the in , and persisted until the stage of the , approximately 90 million years ago. Fossils of carnosaurians have been recovered from every continent except , highlighting their global distribution during periods of peak diversity in the and . Historically, Carnosauria was erected in the as an informal grouping for all large predatory theropods, encompassing a broad array of "carnosaurs" like allosaurids, megalosaurids, and even tyrannosaurids, based on shared traits such as robust skulls and bipedal locomotion. However, cladistic analyses since the late have refined this , establishing Carnosauria as a monophyletic distinct from basal tetanurans like and excluding tyrannosaurids, which are now placed within due to of large size across theropod lineages. Key subgroups include (e.g., from the of ), Carcharodontosauridae (e.g., Carcharodontosaurus and Giganotosaurus from Cretaceous and ), and Sinraptoridae (e.g., Sinraptor from ), each showcasing regional variations in predatory adaptations. Notable for their role as top predators, carnosaurians hunted large herbivores like sauropods and ornithischians, contributing to the dynamic food webs of and terrestrial environments. Carnosaurians are distinguished by several synapomorphies, including robust with serrated, blade-like teeth suited for slashing flesh, large orbits for enhanced , and pneumatic vertebrae that reduced body weight despite their massive sizes—often exceeding 10 meters in length and several tons in . Unlike the reduced forelimbs of tyrannosaurids, many carnosaurians retained relatively strong arms with functional claws, potentially aiding in grasping prey. Modifications in the and hindlimbs, such as a longer than the , supported their bipedal stance and high-speed pursuits, while features like pronounced nuchal crests on the anchored powerful muscles. Their decline by the mid-Cretaceous coincided with the of advanced coelurosaurians and abelisaurids, though carnosaurians like carcharodontosaurids briefly dominated southern continents as the largest known land carnivores. Ongoing discoveries continue to illuminate their evolutionary history, underscoring Carnosauria's importance in understanding theropod diversification and the origins of avian traits.

History of Study

Early Classifications

The term "Carnosauria" was coined by Friedrich von Huene in 1920 to describe a group of large carnivorous saurischians. Earlier, in 1870, classified early known large theropods such as Megalosaurus bucklandii and Poekilopleuron bucklandii within the subgroup of Dinosauria in his paper "On the classification of the Dinosauria," noting their robust builds, serrated teeth suited for tearing flesh, and overall resemblance to modern carnivores. These classifications emphasized large theropods as formidable, bipedal predators from the , distinct from smaller or herbivorous dinosaurs. The discovery of significant fossils played a crucial role in shaping these early groupings. For instance, in 1877, described Allosaurus fragilis based on remains from the in , recognizing it as a large carnivorous that fit within the emerging framework for large theropods due to its powerful jaws and sharp, serrated dentition. By the mid-20th century, Alfred Sherwood Romer broadened Carnosauria considerably in his 1956 monograph Osteology of the Reptiles, treating it as a that incorporated families such as , , and , all unified by their large body sizes, strong limb girdles, and carnivorous dentition. This expansion reflected the era's descriptive taxonomy, which prioritized morphological similarities over evolutionary relationships, leading to inclusions like Tyrannosaurus rex within Carnosauria—a placement that persisted until cladistic analyses in the 1980s reassigned tyrannosaurids to based on shared derived traits such as pneumatic vertebrae and encephalization.

Cladistic Developments

The shift to cladistic methods in the marked a pivotal change in understanding Carnosauria, moving beyond the paraphyletic wastebasket groupings of earlier classifications toward rigorous phylogenetic analyses that emphasized shared derived characters (synapomorphies) and monophyletic clades. Gauthier's seminal 1986 study employed manual cladistic analysis to redefine and its subgroups, positioning Carnosauria as a monophyletic clade within that included taxa such as but explicitly excluded birds (Aves) and tyrannosaurids, which were instead allied with as the to Carnosauria. This redefinition narrowed Carnosauria to basal tetanurans, emphasizing its distinction from more derived avian lineages and highlighting the group's evolutionary position as a major branch of advanced . Building on Gauthier's framework, Roger B. J. Benson's analyses from 2008 to 2010 further refined Carnosauria through expanded datasets and computational methods, confirming it as the sister group to within and incorporating alongside Allosauroidea as core components. Benson's 2008 preliminary phylogeny focused on basal tetanurans, using parsimony-based analyses to resolve relationships among European "megalosaurs" and demonstrate endemism, while his 2010 comprehensive study of integrated over 300 characters across dozens of taxa to solidify Carnosauria's basal position. These works established Carnosauria as a well-supported of large-bodied predators, distinct from the lighter-built coelurosaurs. Key synapomorphies diagnosing Carnosauria during this period included an elongate contributing to a robust and a reduced that reflected adaptations for powerful terrestrial locomotion, features that distinguished the from both ceratosaurs and coelurosaurs. The exclusion of tyrannosauroids from Carnosauria was particularly reinforced by their possession of coelurosaurian traits, such as extensive skeletal pneumatization resulting in hollow, lightweight bones, which contrasted with the denser skeletal construction typical of carnosaurs. This rejection underscored the of large body size in tyrannosauroids and carnosaurs, prioritizing phylogenetic signal from pneumatic features over superficial similarities in predatory morphology. The adoption of during this era, including early parsimony implementations on theropod character matrices, enabled more robust testing of hypotheses and handling of complex datasets, as seen in Benson's use of software to evaluate branch support and alternative topologies. These methods facilitated the integration of new discoveries into evolving cladograms, providing a quantitative foundation that solidified Carnosauria's narrowed scope and influenced subsequent theropod .

Recent Reassessments

In recent years, phylogenetic analyses have increasingly incorporated new fossil discoveries to refine the boundaries and internal structure of Carnosauria within . The 2019 description of Asfaltovenator vialidadi from the Cañadón Asfalto Formation in provided key evidence for resolving long-standing uncertainties in basal tetanuran relationships. This taxon, represented by a partial including cranial and postcranial elements, was positioned as a basal allosauroid in cladistic analyses, supporting the interpretation of as a paraphyletic grade leading toward more derived Allosauroidea and reinforcing the of Carnosauria as encompassing both groups. A 2020 reassessment of isolated theropod teeth from the Upper Bauru Basin in further clarified the diversity and distribution of advanced carnosaurs in . Through morphological, morphometric, and phylogenetic evaluations, the study confirmed the presence of carcharodontosaurid material and highlighted distinctions from other theropod lineages, supporting Neovenatoridae as a distinct family within Allosauroidea separate from core . This work emphasized the biogeographic implications, indicating that neovenatorids persisted in Gondwanan ecosystems alongside abelisauroids during the . The 2025 naming of Tameryraptor markgrafi from the in marked a significant revision of North African carnosaur . Based on re-examination of historical specimens originally attributed to Carcharodontosaurus saharicus, the new genus exhibits unique features such as a prominent nasal crest, leading to its placement outside typical carcharodontosaurines. The associated phylogenetic analysis proposed the Carcharodontosauriformes to unite carcharodontosaurids with their closest relatives, including neovenatorids and basal allosauroids, thereby expanding the conceptual boundaries of advanced carnosaurs and highlighting regional in the African record. Ongoing debates concerning the placement of have been influenced by post-2020 discoveries and expanded datasets. Earlier hypotheses positioned megaraptorans as allosauroids, but studies from 2022 to 2025, incorporating additional skeletal elements from and , have largely favored placement as basal tyrannosauroids within , though debate persists with some analyses suggesting allosauroid affinity citing shared derived traits in the manus and pelvic girdle that align more closely with neovenatorids than with coelurosaurs. Larger phylogenetic matrices, building on Carrano et al.'s framework and updated in 2023 with over 400 characters and 80 taxa, consistently recover Carnosauria as monophyletic within , positioned as the to and emphasizing the clade's role as a diverse radiation of large-bodied predators from the onward.

Anatomy and Morphology

Cranial and Dental Features

Carnosaur skulls are characteristically elongate and narrow, facilitating a lightweight yet robust structure suited to their predatory lifestyle. A defining feature is the large antorbital fenestra, an opening in the skull that can comprise up to 40% of the total skull length in taxa such as Allosaurus, reducing overall cranial mass while housing pneumatic diverticula. This fenestra is bordered by the maxilla, nasal, lacrimal, and jugal bones, with the maxillary antorbital fossa often exceeding 40% of the antorbital cavity's rostrocaudal length in allosauroids. Additionally, prominent sagittal crests extend along the parietals and squamosals, providing extensive attachment sites for jaw adductor muscles; in Carcharodontosaurus, these crests contribute to a skull reaching approximately 1.6 meters in length. Dental morphology in carnosaurs exemplifies ziphodont adaptations, with teeth that are recurved, laterally compressed, and equipped with fine serrations along the mesial and distal carinae. These serrations typically exhibit a of 1–2 per millimeter, enabling efficient slashing of flesh during feeding. The bears a reduced number of teeth compared to more basal theropods, typically 4–5, which are more upright and D-shaped in cross-section to interlock with opposing dentary teeth. In allosauroids, a subnarial gap separates the from the below the external naris, a feature formed by the subnarial processes of the and nasal bones. Sensory structures in carnosaur skulls include large orbits, which occupy a significant portion of the cranial profile and suggest enhanced capabilities. In Allosaurus, the dorsoventrally elongate oval orbits, combined with forward-facing eye positions, yield a binocular estimated at around 20–30 degrees, aiding during hunts.26[517:BVITD]2.0.CO;2)

Postcranial Skeleton

The postcranial skeleton of carnosaurs exhibits adaptations for supporting large body masses while maintaining locomotor efficiency, with overall lengths ranging from 6 to 12 meters and weights between 1 and 7 metric tons across the group. These dimensions vary by taxon, as seen in at approximately 8-9 meters and 2 tons, and approaching 12-13 meters and up to 8 tons. The skeleton features lightweight construction through extensive pneumatization, where invade bones such as vertebrae and long bones, reducing mass without compromising structural integrity—a trait widespread in theropods including carnosaurs. The includes robust characterized by high neural spines, providing enhanced anchorage for epaxial musculature along the neck. These spines contribute to a stiffened vertebral column, supporting the animal's horizontal posture and facilitating balance with the anteriorly positioned during predation. Presacral vertebrae often display pleurocoels and complex internal pneumatic chambers, further lightening the while preserving rigidity for bipedal locomotion. In the pelvic girdle, a diagnostic feature is the triangular pubic boot formed by the distally expanded and conjoined pubes, which expands more than 30% of the pubis length and serves as a key anchorage point for caudal musculature, enhancing power. The pubis itself measures up to 956 mm in large specimens like , with the at 844 mm, forming a robust basin that distributes weight effectively. The hindlimb demonstrates cursorial adaptations, with the femur consistently longer than the tibia in a ratio of approximately 1.2-1.5, as evidenced in Acrocanthosaurus (femur ~128 cm, tibia ~87 cm). This proportion supports estimated top speeds of up to 40 km/h, enabling effective pursuit of prey despite the animals' size. The forelimb is reduced relative to the hindlimb but retains a functional three-fingered manus with subequal digits I-III, each bearing curved claws up to 20 cm in length suited for grappling and subduing prey. In Acrocanthosaurus, the humerus reaches 37 cm, with the full arm (humerus to digit II) spanning about 105 cm, emphasizing its role in close-quarters manipulation rather than primary locomotion.

Soft Tissue Inferences

Evidence from impressions and provides insights into the soft tissues of carnosaurs, revealing a predominantly scaled without feathers in most taxa. A comprehensive review of non-feather integumentary structures in non-avialan theropods highlights skin impressions in the carcharodontosaurid corcovatus, preserving small, polygonal scales on the pes similar to those in modern birds, indicating a non-feathered, scaly covering across much of the body. This scaled , characterized by non-imbricating, tubercular, and polygonal scales, predominates in carnosaurian theropods, with dermal ossifications or osteoderms present in some taxa, such as isolated examples in allosauroids, providing additional armor-like protection. One notable exception within carnosaurs involves potential evidence for filamentous structures in corcovatus. The original description of this taxon notes ulnar tubercles interpreted as possible quill knobs anchoring protofeathers or pennaceous feathers on the forelimb, a feature otherwise known primarily from coelurosaurian theropods. However, this interpretation remains debated, with alternative explanations including pathological overgrowth, vascular foramina, or misidentified muscle scars, as subsequent analyses question their homology to true quill knobs due to their position and morphology. Reconstructions of carnosaurian musculature, based on skeletal attachment sites, indicate substantial retractor power. In Allosaurus fragilis, a representative allosauroid carnosaur, tail vertebrae exhibit deep sulci marking the origin of the caudofemoralis longus muscle, suggesting an exceptionally large and powerful version of this tail-driven femoral retractor, enabling rapid propulsion during locomotion. This muscle, the primary hindlimb retractor in non-avian theropods, likely accounted for a significant portion of the animal's locomotor force, with its size inferred from the expansive scar areas on caudal transverse processes. Inferences about internal organs derive from body cavity proportions and associated trace fossils. Comparative studies of theropod thoracoabdominal partitioning reveal a large in carnosaurs, accommodating extensive viscera including a voluminous gut suited for processing bulky meals. Coprolites attributed to large theropods, such as a massive specimen containing 30-50% fragmented , demonstrate partial of osseous material, implying a capacious intestinal tract with acidic conditions capable of breaking down over extended retention times. Coloration patterns in carnosaurs are inferred through analogy to preserved theropod integuments and modern large predators. While direct melanosome evidence is lacking for carnosaurs, —darker dorsal surfaces grading to lighter ventral areas—appears in related theropods like , suggesting cryptic to reduce visibility against skylines and ground, a pattern common in extant large carnivores for predation. This likely applied to carnosaurs, enhancing concealment in forested or open habitats despite their size.

Systematics and Classification

Definitional Framework

Carnosauria is formally defined in modern cladistic analyses as all theropods more closely related to fragilis than to Neornithes (modern birds), a stem-based definition proposed by Padian et al. (1999) to ensure phylogenetic precision and nomenclatural stability. This definition captures a monophyletic group of large-bodied theropod dinosaurs within , focusing on core allosauroid lineages while excluding more basal or derived theropods, such as spinosaurids. Some analyses have proposed alternative node-based definitions, such as the most inclusive clade containing Allosaurus fragilis and Neovenator salerii, to encompass their last common ancestor and all descendants, emphasizing advanced allosauroids. However, the stem-based approach aligns with consensus efforts to restrict the clade to taxa sharing a closer affinity with allosaurids than to coelurosaurs or other tetanurans, avoiding historical paraphyly where the term encompassed disparate large carnivores. Within Carnosauria, the subclade Allosauroidea comprises and (with Neovenatoridae recognized as a distinct in many analyses), united by synapomorphies such as the presence of a pubic in the , which indicates advanced pneumaticity and structural reinforcement for large body sizes. Exclusion criteria further delineate the group by contrasting it with outgroups like and , both of which possess a (wishbone) that is also present in carnosaurians and thus not diagnostic for exclusion. Post-2000 nomenclatural revisions have prioritized these phylogenetic definitions to maintain stability, deliberately avoiding paraphyletic assemblages that previously broadened Carnosauria to include unrelated megalosaurids or spinosaurids. These efforts reflect a consensus in theropod systematics to align taxonomy with cladistic evidence from comprehensive analyses.

Core Phylogenetic Structure

Carnosauria, equivalent to Allosauroidea in modern usage, is positioned as the sister clade to Coelurosauria within the larger clade Avetheropoda, part of Tetanurae. Basal tetanurans such as those in Megalosauridae and Spinosauridae form a paraphyletic grade outside Carnosauria, leading to the more derived allosauroids. This structure reflects a successive branching pattern in tetanuran evolution. Within Allosauroidea, the primary branches include Allosauridae, encompassing Allosaurus fragilis and Saurophaganax maximus as representative North American Late Jurassic taxa; Carcharodontosauridae, featuring large Cretaceous forms like Giganotosaurus carolinii from South America and Carcharodontosaurus saharicus from Africa; and Neovenatoridae, which includes Neovenator salerii from the Early Cretaceous of Europe and Siats meekerorum from the Early Late Cretaceous of North America. These clades are defined by shared derived traits such as pneumatic vertebrae and specialized cranial features adapted for large-prey predation. Phylogenetic analyses provide strong support for this structure, with Allosauroidea exhibiting Bremer decay indices greater than 3 and bootstrap support exceeding 70% in recent (2025) cladistic analyses incorporating extensive character matrices. Neovenatoridae, in particular, shows high robustness with Bremer support values over 5. Time-calibrated phylogenies place the origin of Carnosauria in the around 174 Ma, coinciding with early tetanuran diversification, while major within Allosauroidea occurred during the . The consensus cladogram can be textually depicted as follows, showing nested relationships:

[Tetanurae](/page/Tetanurae) ├── Basal Tetanurae (e.g., Chuandongocoelurus, [Megalosauridae](/page/Megalosauridae), [Spinosauridae](/page/Spinosauridae)) ├── Avetheropoda │ ├── Carnosauria (Allosauroidea) │ │ ├── [Metriacanthosauridae](/page/Metriacanthosauridae) (basal allosauroids) │ │ ├── [Allosauridae](/page/Allosauridae) (*[Allosaurus](/page/Allosaurus)*, *[Saurophaganax](/page/Saurophaganax)*) │ │ ├── [Carcharodontosauridae](/page/Carcharodontosauridae) (*[Carcharodontosaurus](/page/Carcharodontosaurus)*, *[Giganotosaurus](/page/Giganotosaurus)*) │ │ └── Neovenatoridae (*[Neovenator](/page/Neovenator)*, *[Siats](/page/Siats)*) │ └── [Coelurosauria](/page/Coelurosauria)

[Tetanurae](/page/Tetanurae) ├── Basal Tetanurae (e.g., Chuandongocoelurus, [Megalosauridae](/page/Megalosauridae), [Spinosauridae](/page/Spinosauridae)) ├── Avetheropoda │ ├── Carnosauria (Allosauroidea) │ │ ├── [Metriacanthosauridae](/page/Metriacanthosauridae) (basal allosauroids) │ │ ├── [Allosauridae](/page/Allosauridae) (*[Allosaurus](/page/Allosaurus)*, *[Saurophaganax](/page/Saurophaganax)*) │ │ ├── [Carcharodontosauridae](/page/Carcharodontosauridae) (*[Carcharodontosaurus](/page/Carcharodontosaurus)*, *[Giganotosaurus](/page/Giganotosaurus)*) │ │ └── Neovenatoridae (*[Neovenator](/page/Neovenator)*, *[Siats](/page/Siats)*) │ └── [Coelurosauria](/page/Coelurosauria)

This nesting highlights the progressive specialization from basal tetanurans to advanced allosauroids.

Debated Inclusions and Alternatives

The phylogenetic placement of has been highly debated, with early analyses from 2012 positioning it as a basal coelurosaur clade characterized by large manual claws and robust forelimbs. Subsequent studies between 2016 and 2025 shifted toward an allosauroid affinity, recovering within Neovenatoridae due to shared features such as pneumatic vertebrae and elongated manual unguals, exemplified by the reanalysis of as a derived carcharodontosaurian. This realignment highlights the 's Gondwanan distribution and survival into the , though some analyses still favor coelurosaurian ties based on cranial proportions. The status of remains contentious, often interpreted as a paraphyletic grade rather than a monophyletic group. The 2019 description of Asfaltovenator vialidadi from the of supported a monophyletic Carnosauria encompassing Allosauroidea and , but positioned megalosauroid subclades (e.g., , , ) as sequential outgroups to Allosauroidea, underscoring extensive in early tetanuran evolution. This configuration implies represents a basal grade leading to more derived allosauroids, challenging prior views of its unity. Alternative taxonomic hypotheses have proposed an expanded Carnosauria incorporating spinosauroids alongside allosauroids and megalosauroids, based on shared predatory adaptations and origins. However, such broad definitions have been largely rejected in favor of a restricted Carnosauria excluding , as evidenced by consistent recoveries in large-scale matrices emphasizing dental and postcranial distinctions. Recent discoveries like Tameryraptor markgrafi from the of further refine carnosaur , elevating the prominence of Carcharodontosauriformes within Allosauroidea. This new carcharodontosaurid, distinguished by a modest nasal horn and symmetrical maxillary teeth, suggests greater diversity among North African forms and potential exclusion of some basal allosauroids from core carcharodontosaurid clades due to biogeographic barriers like the Trans-Saharan seaway. Its phylogeny reinforces Carcharodontosauridae's roots while highlighting faunal provincialism in the . Methodological challenges in carnosaur phylogeny often stem from the sensitivity of analyses to character selection and scoring, particularly in ambiguous traits like manual morphology and phalangeal counts. Variations in coding hand claw features—such as the number and curvature of unguals—can shift placements of peripheral taxa like megaraptorans between allosauroids and coelurosaurs, as demonstrated in comparative matrix evaluations. This instability underscores the need for robust, multi-character datasets to resolve ongoing debates.

Paleobiology and Ecology

Predatory Adaptations

Carnosaurians exhibited predatory adaptations centered on a combination of cranial mechanics and locomotor capabilities that favored slashing and tearing over sustained crushing bites. Biomechanical models of the Allosaurus jaw indicate bite force estimates ranging from approximately 1 to 3 kN at the tooth row, significantly lower than in later tyrannosaurids, enabling rapid, slashing attacks to inflict deep wounds rather than bone-crushing grips. This strategy is supported by the lightweight skull construction and recurved, serrated teeth, which facilitated puncturing and pulling flesh from large prey. The denticles on these teeth enhanced tissue damage by creating irregular tears during withdrawal, optimizing efficiency in dismembering carcasses or debilitating live targets. Evidence from fossil trackways suggests carnosaurians employed a pursuit or hunting style, leveraging bipedal speed and powerful hindlimbs for short bursts of . Track sites preserving theropod gaits indicate velocities of 20-30 km/h for individuals comparable in size to , allowing them to close distances on slower herbivores in open terrains. Large, curved claws on the manus and pes further aided in and subduing prey during close encounters, complementing their role as active hunters rather than obligate . Prey preferences among carnosaurians targeted massive herbivores, as evidenced by theropod bite traces on sauropod bones from the . Allosaurus-inflicted marks, including deep punctures and grooves, appear on diplodocid elements such as caudal vertebrae, indicating attacks on vital areas to cause hemorrhage or immobilize subadult individuals. These traces, often unhealed and clustered on low-meat-yield bones, suggest opportunistic feeding on weakened or juvenile sauropods, which dominated Morrison ecosystems. Sensory adaptations enhanced detection and targeting of prey, with enlarged external nares and cavities in allosauroid skulls pointing to a keen for tracking over distances. Combined with proportionally large orbits—up to 20% of skull length in —this implies high visual acuity for spotting movement in diurnal settings, though binocular overlap was limited to about 20 degrees due to laterally positioned eyes. In the Early to mid-Cretaceous, carcharodontosaurids occupied niches across Gondwanan landmasses, preying on titanosaurian sauropods in what are now and . Fossils from deposits in , such as those of , alongside South American taxa like , confirm their dominance as the largest terrestrial carnivores, with body masses exceeding 6 tons enabling predation on herbivores over 50 tons. This partitioning underscores their ecological role in structuring food webs before the rise of abelisaurids and tyrannosaurids.

Growth, Pathology, and Behavior

Bone histology studies of carnosaurians, particularly , reveal rapid juvenile growth rates characteristic of large theropods, with maximum somatic growth occurring at approximately 15 years of age, when annual body mass increase peaked at 148 kg. This rapid phase allowed individuals to attain substantial size early in , with the largest sampled specimens estimated to be 13–19 years old and skeletal maturity reached between 22 and 28 years. Such growth patterns, inferred from lines of arrested growth and fibrolamellar tissue, indicate a determinate growth similar to that of modern large reptiles but accelerated to support a predatory lifestyle. Pathological evidence from carnosaurian fossils demonstrates a high frequency of traumatic injuries, including healed fractures and bite-induced infections, likely resulting from intraspecific aggression or unsuccessful hunts. For instance, the subadult specimen MOR 693 preserves at least 19 distinct injuries across its skeleton, such as broken ribs, a fractured , and an infected toe , many of which show signs of healing and reinjury. These pathologies are common in large-bodied theropods, with multiple lesions per individual underscoring the physical demands of predation and potential combat over resources or mates. The presence of localized infections from deep bite wounds further suggests that affected individuals survived without extensive , consistent with solitary or small-group living rather than large packs. Behavioral inferences from indicate that carnosaurians were primarily active predators but engaged in scavenging when opportunities arose, as evidenced by theropod bite marks on lower-economy bones (e.g., ribs and vertebrae) from the Mygatt-Moore Quarry, where marks are concentrated on less nutritious elements typical of post-mortem feeding. Although direct gastric contents are rare, the pattern of bone modification supports occasional scavenging supplementing hunting, particularly in stressed ecosystems. Limb bone robusticity and orbit morphology in large carnosaurians like suggest diurnal activity patterns, contrasting with the nocturnal habits of smaller theropods and aligning with visual adaptations for daytime predation.

Reproductive and Ontogenetic Insights

Putative medullary bone, a tissue associated with egg-laying, has been tentatively reported in histological analyses of Allosaurus specimens, though its identification remains uncertain and debated. This tissue has been confirmed in other non-avian theropods such as Tyrannosaurus. However, no direct fossil evidence of nests or eggs has been attributed to carnosaur taxa, limiting insights to indirect indicators from related theropods. Ontogenetic studies of demonstrate significant morphological changes during development, including proportional shifts in cranial features. Morphometric analyses of skull elements reveal bimodal distributions along allometric trajectories, interpreted as potential , with clusters possibly representing male and female variants in robusticity and size. Juveniles exhibit proportionally larger heads relative to body size compared to adults, a pattern consistent with early ontogenetic stages in theropods where enlarged crania may facilitate prey capture or sensory functions. These changes reflect a dynamic growth strategy transitioning from juvenile to adult forms. Bonebeds provide key evidence for social behavior in carnosaurs, particularly in , where disarticulated remains of at least seven to nine individuals from a single locality represent a range of growth stages, from juveniles to adults. This assemblage suggests gregarious habits, potentially forming packs that enabled cooperative hunting of large prey, as inferred from the spatial association and ontogenetic diversity. Growth trajectories in carnosaurs follow sigmoid (S-shaped) curves, as reconstructed from histology and length data in , with rapid juvenile growth peaking around 15 years of age before asymptotic stabilization. Maturity is estimated to occur between 20 and 25 years, accompanied by patterns typical of dinosaurs, where external fundamental osteons indicate continued, albeit slower, skeletal deposition into adulthood. The mixed-age composition of the bonebed implies possible or age-segregated social structures, analogous to those in crocodilians where juveniles remain near adults for . Such assemblages suggest that carnosaurs may have exhibited dynamics, with size segregation facilitating resource partitioning or group defense during vulnerable developmental phases.

Paleobiogeography and Distribution

Temporal Range

The temporal range of Carnosauria encompasses the to the , approximately 168 to 92 million years ago (Ma). The earliest definitive records appear in the , during the stage (~168 Ma), represented by basal members such as jiangi from the in , , which provides evidence of early diversification among large-bodied theropods in . Carnosauria reached its peak diversification in the (Kimmeridgian to stages, 163–145 Ma), particularly within , where dominated as apex predators; notable examples include Allosaurus fragilis from the in , a key biostratigraphic unit spanning the late Kimmeridgian to early . In the , carnosaurian lineages persisted primarily through during the (Berriasian to stages, 145–100 Ma), with widespread occurrences across and , but underwent a marked decline after ~100 Ma as tyrannosauroids and abelisauroids rose to prominence. The final known occurrences date to around 92 Ma in the stage, exemplified by Shaochilong maortuensis from the Ulansuhai Formation () in , , marking the effective end of the group as it was ecologically displaced. Biostratigraphic correlations highlight key formations such as the Morrison (late –, ~155–147 Ma) for Late Jurassic allosaurids and the Bahariya Formation (early , ~100–95 Ma) for North African carcharodontosaurids like Carcharodontosaurus saharicus, underscoring the group's temporal progression across major depositional basins.

Geographic Patterns and Dispersal

Carnosauria, encompassing large theropod dinosaurs such as allosauroids, originated in during the , with significant fossil records from and . In , Allosaurus fragilis dominated the , a vast depositional basin spanning multiple states, where it represented the in diverse ecosystems. European occurrences include Allosaurus material from Portugal's Lourinhã Formation, indicating early transatlantic dispersal or shared ancestry across the proto-Atlantic rift. The group underwent a notable radiation in during the Early to mid-Cretaceous, particularly in and , where carcharodontosaurids became prominent large predators. In , Giganotosaurus carolinii from Argentina's exemplifies this diversification, dating to approximately 99 million years ago in the stage. African records, such as Carcharodontosaurus from North African deposits, further highlight this southern expansion, with taxa adapted to arid, riverine environments. Asian records of Carnosauria remain limited, primarily consisting of isolated or partial specimens from the mid-Cretaceous. , a carcharodontosaurid from China's Ulansuhai Formation ( stage), represents one of the few definitive Asian members, suggesting possible vicariance between Laurasian and Gondwanan lineages following continental fragmentation. Dispersal events for Carnosauria likely occurred in the Mid-Jurassic via land bridges across the still-intact supercontinent, allowing initial global spread from Laurasian origins. Post- breakup, progressive isolation of Gondwanan landmasses from the onward promoted endemic radiations, such as the diversification of carcharodontosaurids in southern continents, as evidenced by vicariance patterns in theropod distributions. Biogeographic patterns of Carnosauria delineate at least four major provinces: a Laurasian Jurassic province centered on and , a Gondwanan Cretaceous province dominant in and , and sparse records in and polar regions. Australia and exhibit particularly low diversity, with only recent discoveries of carcharodontosaurian fossils in Australia's strata confirming limited presence, likely due to high-latitude barriers and isolation; in 2025, the first carcharodontosaur fossils were reported from the ~120 Ma Strzelecki Group in Victoria.

References

  1. https://www.geol.umd.edu/~tholtz/gaiaphyl.pdf
  2. https://ucmp.berkeley.edu/diapsids/saurischia/theropoda.html
  3. https://www.nature.com/articles/s41598-019-53672-7
  4. https://theropoddatabase.github.io/Carnosauria.htm
  5. https://eofauna.com/book/theropoda_records_refB.pdf
  6. http://www.si.edu/object/nmnhpaleobiology_3451264
  7. https://upload.wikimedia.org/wikipedia/commons/a/a9/The_Osteology_of_the_Reptiles.pdf
  8. https://theropoddatabase.github.io/Tyrannosauroidea.html
  9. https://archive.org/download/biostor-110202/biostor-110202.pdf
  10. https://www.researchgate.net/publication/230808558_The_phylogeny_of_Tetanurae_Dinosauria_Theropoda
  11. https://royalsocietypublishing.org/doi/10.1098/rsos.242238
  12. http://www-personal.umich.edu/~wilsonja/JAW/Publications_files/Sereno%2526al1996.pdf
  13. https://doc.rero.ch/record/14265/files/PAL_E1381.pdf
  14. https://pmc.ncbi.nlm.nih.gov/articles/PMC6984342/
  15. https://ix.cs.uoregon.edu/~kent/paleontology/binocularVision/index.html
  16. https://www.researchgate.net/publication/382704993_Gigantism_in_Theropods_A_Study_of_Giganotosaurus_carolinii_and_Its_Ecological_Impact
  17. https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/g2000n2a3.pdf
  18. https://ucmp.berkeley.edu/diapsids/saurischia/carnosauria.html
  19. https://onlinelibrary.wiley.com/doi/abs/10.1111/brv.12829
  20. https://www.researchgate.net/publication/317151437_Origin_attachments_of_the_caudofemoralis_longus_muscle_in_the_Jurassic_dinosaur_Allosaurus
  21. https://www.researchgate.net/publication/232796267_A_king-size_theropod_coprolite
  22. https://www.sciencedirect.com/science/article/pii/S0960982217311971
  23. https://www.tandfonline.com/doi/abs/10.1080/02724639909506978
  24. https://link.springer.com/article/10.1007/s00114-009-0614-x
  25. https://www.tandfonline.com/doi/abs/10.1080/14772019.2011.630927
  26. https://doi.org/10.1371/journal.pone.0311096
  27. https://www.research.ed.ac.uk/files/8232966/PDF_BrusatteSereno2008AllosauroidPhylogeny.pdf
  28. https://www.researchgate.net/publication/12097920_The_bite_of_Allosaurus
  29. https://anatomypubs.onlinelibrary.wiley.com/doi/10.1002/ar.20977
  30. https://www.sciencedirect.com/science/article/pii/S0960982218303713
  31. https://pmc.ncbi.nlm.nih.gov/articles/PMC9285543/
  32. https://pmc.ncbi.nlm.nih.gov/articles/PMC10655710/
  33. https://svpow.com/2023/11/14/new-paper-theropod-bite-marks-on-morrison-sauropod-bones/
  34. https://people.ohio.edu/witmerl/Downloads/2011_Zelenitsky_et_al._theropod_olfaction.pdf
  35. https://www.tandfonline.com/doi/full/10.1080/02724634.2023.2295518
  36. https://www.researchgate.net/publication/228671730_Binocular_vision_in_theropod_dinosaurs
  37. https://www.nature.com/articles/ncomms3827
  38. https://www.nature.com/articles/s41598-022-09272-z
  39. https://onlinelibrary.wiley.com/doi/10.1002/jmor.10406
  40. https://www.frontiersin.org/journals/earth-science/articles/10.3389/feart.2021.641060/full
  41. https://peerj.com/articles/940/
  42. https://pmc.ncbi.nlm.nih.gov/articles/PMC4435507/
  43. https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0233115
  44. https://pubmed.ncbi.nlm.nih.gov/21493820/
  45. https://pmc.ncbi.nlm.nih.gov/articles/PMC7017430/
  46. https://www.nature.com/articles/s41467-018-07621-z
  47. https://www.researchgate.net/publication/272151969_A_morphometric_analysis_of_Allosaurus
  48. https://royalsocietypublishing.org/doi/10.1098/rspb.2005.3071
  49. https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/g2006n1a4.pdf
  50. https://www.newscientist.com/article/dn9007-giant-dino-predators-may-have-hunted-in-packs/
  51. https://pubmed.ncbi.nlm.nih.gov/16380967/
  52. https://www.researchgate.net/publication/233708008_A_distinct_dinosaur_life_history
  53. https://www.researchgate.net/publication/236855910_The_age_of_the_Morrison_Formation
  54. https://www.cell.com/current-biology/fulltext/S0960-9822(22)00860-0
  55. https://mapress.com/zt/article/view/zootaxa.2334.1.1
  56. https://www.sciencedirect.com/science/article/pii/S0037073806001357
  57. https://nhmu.utah.edu/allosaurus-fragilis
  58. https://pubs.geoscienceworld.org/jgs/article-lookup?doi=10.1144/gsjgs.156.3.0449
  59. https://www.tandfonline.com/doi/abs/10.1080/02724634.1998.10011027
  60. https://www.scielo.br/j/paz/a/H8rTsTK5cF8Lqyz664GxgTP/?format=html&lang=en
  61. https://royalsocietypublishing.org/doi/pdf/10.1098/rspb.2001.1921
  62. https://www.monash.edu/science/news-events/news/2025/australias-first-carcharodontosaur-fossils-unearthed-along-victorias-cretaceous-coast
Add your contribution
Related Hubs
User Avatar
No comments yet.