Eocarcharia Dinops
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''Eocarcharia'' (meaning "dawn shark") is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaurs found in what is now the western
Ténéré Desert The Ténéré (Tuareg: Tenere, literally: "desert") is a desert region in south central Sahara. It comprises a vast plain of sand stretching from northeastern Niger to western Chad, occupying an area of over . The Ténéré's boundaries are said ...
of Niger. It is known from several skull bones collected in 2000 by an expedition to the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
(
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...
ages)
Elrhaz Formation The Elrhaz Formation is a geological Formation (geology), formation in Niger, West Africa. Its strata date back to the Early Cretaceous, about 125 to 112 million years ago. Dinosaur remains are among the fossils that have been recovered from the ...
(Gadoufaoua locality) led by American paleontologist
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago who has discovered several new dinosaur species on several continents, including at sites in Inner Mongolia, Argentina, Morocco and Niger. ...
. They were then described in 2008 by Sereno and
Steve Brusatte Stephen Louis "Steve" Brusatte FRSE (born April 24, 1984) is an American author and evolutionary biologist who specializes in the anatomy and evolution of dinosaurs. He was educated at the University of Chicago for his Bachelor's degree, at the ...
. The genus contains a single species, ''Eocarcharia dinops''. Sereno and Brusatte identified the remains as a new
carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
, but later studies suggested that the species is chimaeric, comprising bones of multiple unrelated taxa. Some of the ''Eocarcharia'' material, including the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
(name-bearing) specimen, likely belongs to a
baryonychine Baryonychinae is an extinct clade or subfamily of spinosaurids from the Early Cretaceous of Europe and West Africa. The clade was named by Charig & Milner in 1986 and defined by Sereno et al. in 1998 and Holtz et al. in 2004 as all taxa more close ...
spinosaurid Spinosauridae (or spinosaurids) is a clade or Family (taxonomy), family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. Spinosaurid fossils have been recovered worldwide, including Africa, Europe, South America, and Asia. ...
. This would render ''Eocarcharia'' a member of this group, closely related to the coeval ''
Suchomimus ''Suchomimus'', from Ancient Greek ''σούχος'' (''soúkhos''), meaning "crocodile", and Latin ''mimus'', meaning "actor", is a genus of spinosaurid dinosaur that lived between 125 and 112 million years ago in what is now Niger, North A ...
''. Meanwhile, the definitively carcharodontosaurid bones, a and teeth, belong to a separate distinct taxon. Little is known about ''Eocarcharia'' due to its fragmentary and chimeric nature. When considered a carcharodontosaurid, it was estimated to be long, making it smaller than derived carcharodontosaurids like ''
Giganotosaurus ''Giganotosaurus'' ( ) is a genus of large theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period (geology), period, approximately 99.6 to 95 million years ago. The holotype specim ...
,'' and ''
Carcharodontosaurus ''Carcharodontosaurus'' (; ) is a genus of large carnivorous theropod dinosaur that lived in Northwest Africa from about 100 to 94 million years ago during the Cenomanian age of the Cretaceous. Two teeth of the genus, now lost, were first des ...
''. The (bone behind the orbit) is robust with a large brow, a distinct characteristic of ''Eocarcharia''. It was possibly covered in
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
and used for
head-butting A headbutt or butt is a targeted strike with the head, typically involving the use of robust parts of the headbutter's cranium as the area of impact. The most effective headbutts strike the most sensitive areas of an opponent, such as the nose, ...
with other individuals of its species. ''Eocarcharia'' lived in an environment with rivers and vast
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high Discharge (hydrolog ...
s alongside many other dinosaurs,
pterosaurs Pterosaurs are an extinct clade of flying reptiles in the Order (biology), order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosau ...
,
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. Extinct crocodylomorphs were considerably more ...
, and freshwater animals.


Discovery and naming

In 2000, American paleontologist
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago who has discovered several new dinosaur species on several continents, including at sites in Inner Mongolia, Argentina, Morocco and Niger. ...
led an expedition conducted with the
University of Chicago The University of Chicago (UChicago, Chicago, or UChi) is a Private university, private research university in Chicago, Illinois, United States. Its main campus is in the Hyde Park, Chicago, Hyde Park neighborhood on Chicago's South Side, Chic ...
and funded by the
National Geographic Society The National Geographic Society, headquartered in Washington, D.C., United States, is one of the largest nonprofit scientific and educational organizations in the world. Founded in 1888, its interests include geography, archaeology, natural sc ...
. The purpose of this trip was to explore fossiliferous
sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
outcrops in a site on the western edge of the Nigerian
Ténéré Desert The Ténéré (Tuareg: Tenere, literally: "desert") is a desert region in south central Sahara. It comprises a vast plain of sand stretching from northeastern Niger to western Chad, occupying an area of over . The Ténéré's boundaries are said ...
known as Gadoufaoua. These rock layers belong to the
Elrhaz Formation The Elrhaz Formation is a geological Formation (geology), formation in Niger, West Africa. Its strata date back to the Early Cretaceous, about 125 to 112 million years ago. Dinosaur remains are among the fossils that have been recovered from the ...
, dating to the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
. During the expedition, several isolated theropod skull bones were collected, including a
maxilla In vertebrates, the maxilla (: maxillae ) 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. The two maxil ...
(main tooth-bearing bone of the upper jaw) and two maxilla fragments, five (bone behind the orbit), two —one articulated with the left and one with a partial ( bones)— and five
teeth A tooth (: teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
. These remains were then transported to the University of Chicago for study and preparation before being returned to the Musee National du Niger and deposited under the catalog numbers MNN GAD2-14. In 2008, Sereno and
Steve Brusatte Stephen Louis "Steve" Brusatte FRSE (born April 24, 1984) is an American author and evolutionary biologist who specializes in the anatomy and evolution of dinosaurs. He was educated at the University of Chicago for his Bachelor's degree, at the ...
described all of the remains as belonging to a new genus and species of carcharodontosaurid, named ''Eocarcharia dinops''. They established one of the postorbitals, MNN-GAD2, as the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
(name-bearing) specimen. The generic name, ''Eocarcharia'', derives from the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
words , meaning , and , meaning , referencing the early-diverging nature of ''Eocarcharia'' in relation to its 'shark-toothed' relatives. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
, , is a Greek term meaning "fierce-eyed", referring to the postorbital ornamentation above the eye.


Description

Based on the total fusion of the skull bones referred to ''Eocarcharia'', Sereno & Brusatte identified them as having come from fully mature adult individuals. The only remains confidently referrable to ''Eocarcharia'' consist of the holotype postorbital, four referred postorbitals, a left frontal, and a left prefrontal. Referencing the more complete skulls and skeletons of other carcharodontosaurids like ''
Acrocanthosaurus ''Acrocanthosaurus'' ( ; ) is a genus of carcharodontosaurid dinosaurs 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, ...
'' and ''
Carcharodontosaurus ''Carcharodontosaurus'' (; ) is a genus of large carnivorous theropod dinosaur that lived in Northwest Africa from about 100 to 94 million years ago during the Cenomanian age of the Cretaceous. Two teeth of the genus, now lost, were first des ...
'', they determined that ''Eocarcharia'' was likely about long, half the linear size of the largest, more derived members of the family. Similar estimates were supported by later publications.


Posterior skull

The postorbital bears a thick, robust brow that, likely due to its robusticity, preserves well. This brow is composed of two segments; an anterior (front) portion with boxy dimensions and a posterior (back) portion that forms an ovate orbital boss (a large, thick bony mass). This orbital boss is positioned above the posterodorsal (top back) edge of the
orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
, a trait noted by Sereno and Brusatte to be
diagnostic Diagnosis (: diagnoses) is the identification of the nature and cause of a certain phenomenon. Diagnosis is used in a lot of different academic discipline, disciplines, with variations in the use of logic, analytics, and experience, to determine " ...
(a distinct feature of a species). However, the occurrence of an exaggerated orbital boss is also found in
spinosaurids Spinosauridae (or spinosaurids) is a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. Spinosaurid fossils have been recovered worldwide, including Africa, Europe, South America, and Asia. Their remains have ...
. An orbital boss is also found in ''
Mapusaurus ''Mapusaurus'' ( Earth lizard) is a genus of giant carcharodontosaurid carnosaurian dinosaur that lived in Argentina during the Turonian age of the Late Cretaceous. Discovery ''Mapusaurus'' was excavated between 1997 and 2001, by the Argen ...
'' and ''
Giganotosaurus ''Giganotosaurus'' ( ) is a genus of large theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period (geology), period, approximately 99.6 to 95 million years ago. The holotype specim ...
'', though the orbital bosses of these genera are
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
as a distinct body from the rest of the postorbital. In contrast, the postorbital bosses of ''Carcharodontosaurus'' and ''Eocarcharia'' lack the extensive ossification seen in ''Mapusaurus'' and ''Giganotosaurus''. The margins of the postorbital make up segments of the orbit edges, the , and the . Several of the postorbitals known from ''Eocarcharia'' preserve their articular surfaces (part of the postorbital that articulates with the lacrimal), which are deep and long. These articular surfaces are very small and deep proportionally for a carcharodontosaurid, making this characteristic a diagnostic trait of ''Eocarcharia''. In contrast, the point of contact between the postorbital and the frontal is rugose and bears a unique, plate-shaped
process A process is a series or set of activities that interact to produce a result; it may occur once-only or be recurrent or periodic. Things called a process include: Business and management * Business process, activities that produce a specific s ...
(projection of bone) that interlocks with a concavity on the frontal. Another diagnostic trait of ''Eocarcharia'' is that its bears a narrow facet which articulates the postorbital with the (cheekbone), whereas carcharodontosaurids like ''Carcharodontosaurus'' have broader facets. Midshafts of the
ventral Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
(bottom) ramus (a branch of bone) in spinosaurids and megalosaurids are typically U-shaped in
cross-section Cross section may refer to: * Cross section (geometry) ** Cross-sectional views in architecture and engineering 3D * Cross section (geology) * Cross section (electronics) * Radar cross section, measure of detectability * Cross section (physics) ...
, whereas in ''Eocarcharia'' it is subtriangular in cross-section. An infraoribital process is also present on the ventral ramus of ''Eocarcharia'', however it is distinct from those of other carcharodontosaurids in that it is small, distally (away from body core)-positioned, and rugose. The frontal is known from ''Eocarcharia'', which is proportionally very broad at its mid length, similar to that of ''Carcharodontosaurus''. In dorsal view, the supratemporal (a depression along the
temporal fenestra Temporal fenestrae are openings in the temporal region of the skull of some amniotes, behind the orbit (eye socket). These openings have historically been used to track the evolution and affinities of reptiles. Temporal fenestrae are commonly (al ...
) is greatly exposed, another diagnostic characteristic. In ventral view, the anterior portion of the frontal is exposed and forms the roof of the
olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
section of the
endocranium The endocranium in comparative anatomy is a part of the skull base in vertebrates and it represents the basal, inner part of the cranium. The term is also applied to the outer layer of the dura mater in human anatomy. Structure Structurally, t ...
(part of skull that contains the brain). This portion of the frontal is narrow, in contrast to those of
tetanurans Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, and coelurosaurs (which includes tyrannosauroids, ornithomimosaurs, compsognathids and maniraptorans, ...
which are typically broad. Notably, a prefrontal is also preserved. Independent prefrontals are absent in advanced carcharodontosaurids as they are typically coosified with the lacrimals into a single element, whereas in genera like ''Acrocanthosaurus, Eocarcharia,'' and ''
Concavenator ''Concavenator'' (meaning Cuenca hunter) is a genus of carcharodontosaurian dinosaur that lived in Spain during the Early Cretaceous epoch, about 125 million years ago. The genus contains a single species, ''Concavenator corcovatus'', named and ...
'', they are unfused. The prefrontal bears an extended process engineered to articulate with the deep, squared articular notch on the frontal. On the ventral (bottom) side of the frontal, theropods typically feature an extended process. that travels along the posteromedial (back inner) portion of the bone. ''Eocarcharia'' lacks this characteristic and is instead enlarged, distinct, and lacks the ossification.


Referred maxilla and teeth

One nearly complete maxilla and two central maxilla portions were initially referred to ''Eocarcharia'' and definitively belong to a carcharodontosaurid or closely allied taxon. The dentary has 15
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * M ...
(tooth sockets). It is generally flat, which would have given this taxon a relatively narrow snout. (small pits and grooves) cover the external surface. The , a depression on the side of the maxilla, is pierced by three
fenestrae A fenestra (fenestration; : fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biology, biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomy, ...
(holes), the largest of which is the subtriangular promaxillary fenestra, followed by the maxillary fenestra, which is subequal in size. A smaller round accessory fenestra is present above these two. Most of the alveoli in the complete maxilla preserve
replacement teeth Replacement(s) or Replace may refer to: Music * The Replacements (band), an American alternative rock band Film and television * ''The Replacements'' (film), a 2000 American sports comedy * ''The Replacement'' (2021 film), a 2021 Spanish thr ...
. The are transversely compressed. Unlike many carcharodontosaurid teeth, those referred to ''Eocarcharia'' are not blade-shaped and they lack straight posterior (cutting edges on the back of the crown) and enamel wrinkles with high relief.


Classification


As a carcharodontosaurid

In their
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
, Sereno & Brusatte (2008) recognized ''Eocarcharia'' as an allosauroid theropod in an early-diverging position within the family Carcharodontosauridae. They noted that the resolution of their trees was more stable without ''Eocarcharia'', since it could not be scored for most of the characters in their dataset. Regardless, their strict consensus tree recovered it as the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to the North American ''Acrocanthosaurus''.In the same 2008 paper, Sereno and Brusatte named '' Kryptops palaios'' on the basis of an
abelisaurid Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are foun ...
maxilla and postcranial remains. They assigned the postcranial material to the same individual as the maxilla based on their close association and alleged basal abelisaurid features in the vertebrae and pelvis. In 2012, Matthew Carrano and colleagues considered ''Kryptops palaios'' to be a
chimera Chimera, Chimaera, or Chimaira (Greek for " she-goat") originally referred to: * Chimera (mythology), a fire-breathing monster of ancient Lycia said to combine parts from multiple animals * Mount Chimaera, a fire-spewing region of Lycia or Cilicia ...
(specimen composed of multiple species), and stated that its postcranial remains, especially the pelvis and sacrum, may actually belong to a carcharodontosaurid, possibly ''Eocarcharia''. However, these bones do not overlap with the ''Eocarcharia'' holotype, which consists only of an isolated postorbital. This hypothesis was supported by later studies, who agreed that the postcranial remains belonged to an
allosauroid 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 Carn ...
, carcharodontosaurid, or a
metriacanthosaurid Metriacanthosauridae (Greek for "moderately-spined lizards") is an extinct family of allosauroid theropod dinosaurs that lived in Europe and Asia from the Middle Jurassic to the Early Cretaceous. The family is split into two subgroups: Metriacant ...
. In their 2024 description of a new specimen of the giant carcharodontosaurid ''
Taurovenator ''Taurovenator'', from Latin ''taurus'', meaning "bull", and ''venator'', meaning "hunter", is an extinct genus of large carcharodontosaurid theropod dinosaurs that lived in what is now Argentina during the Cenomanian age of the Late Cretaceous, ...
'', Rolando and colleagues recovered ''Eocarcharia'' as an early-diverging member of the clade, branching after ''
Neovenator ''Neovenator'' ( nˈiːə͡ʊvˌɛne͡ɪtə; "new hunter") is a genus of carcharodontosaurian theropod dinosaur. It is known primarily from several skeletons found in the Early Cretaceous (Hauterivian-Barremian) Wessex Formation on the south co ...
''. The results of their
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
are displayed in the
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
below: The skull bones of ''Eocarcharia'' were not found in association and they belong to multiple individuals. As such, the taxonomic identity of these bones—and whether or not they can all be referred to the same taxon—has been debated. In their description of the carcharodontosaurid ''
Tameryraptor ''Tameryraptor'' ("thief from the beloved land") is an extinct genus of large carcharodontosaurid dinosaur that lived during the Late Cretaceous (Cenomanian age) in what is now Egypt. It is known from a partial skeleton collected in rock layers ...
'', Kellermann, Cuesta & Rauhut (2025) scored the holotype postorbital (including a referred frontal with articular surfaces that match those of the postorbital) and referred maxilla as separate
operational taxonomic unit An operational taxonomic unit (OTU) is an operational definition used to classify groups of closely related individuals. The term was originally introduced in 1963 by Robert R. Sokal and Peter H. A. Sneath in the context of numerical taxonomy, wh ...
s (OTUs) to test the likelihood that they belonged to the same taxon. In their analyses, both OTUs were consistently recovered in different positions, supporting their status as distinct taxa. The maxilla was reliably recovered as a carcharodontosauriform, either as a non- carcharodontosaurine carcharodontosaurid or as a
metriacanthosaurid Metriacanthosauridae (Greek for "moderately-spined lizards") is an extinct family of allosauroid theropod dinosaurs that lived in Europe and Asia from the Middle Jurassic to the Early Cretaceous. The family is split into two subgroups: Metriacant ...
. On the other hand, the holotype was recovered in positions as a basal carcharodontosaur, or variably inside or outside of Carcharodontosauriformes. They interpreted these results as indicative of the referred maxilla skewing past analyses toward carcharodontosaur affinities for the taxon.


As a spinosaurid

In 2024,
Andrea Cau Andrea Cau is an Italian vertebrate paleontologist. He specialises in the study of dinosaur cladistics. Cau named the unique dromaeosaurid theropod '' Halszkaraptor'' in 2017. He also reanalysed the theropod ''Balaur'', placing it as a basal avial ...
published the results of a comprehensive theropod phylogenetic framework. The following year, Cau and Paterna used an updated version of this dataset to analyze the relationships of Cretaceous theropods from Africa, especially those known from multiple specimens with minimal overlapping material. The researchers scored two separate OTUs for ''Eocarcharia'': the referred maxilla and another comprising the skull roof material (including the referred frontal and prefrontal, and the holotype postorbital). As expected, the maxilla was found to have close affinities with early-diverging carcharodontosaurids. More surprisingly, the skull roof was recovered as the sister taxon to the coeval ''Suchomimus'' as a
baryonychine Baryonychinae is an extinct clade or subfamily of spinosaurids from the Early Cretaceous of Europe and West Africa. The clade was named by Charig & Milner in 1986 and defined by Sereno et al. in 1998 and Holtz et al. in 2004 as all taxa more close ...
within the Spinosauridae. These results were supported by the presence of at least seven shared traits in ''Eocarcharia'' and spinosaurids that are absent in carcharodontosaurs. Cau and Paterna further recognized that two of the
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...
(unique derived traits) proposed for ''Eocarcharia'' by Sereno and Brusatte are also uniquely shared with ''Ceratosuchops'', a baryonychine from the
Wessex Formation The Wessex Formation is a fossil-rich England, English geological formation that dates from the Berriasian to Barremian Stage (stratigraphy), stages of the Early Cretaceous. It forms part of the Wealden Group and underlies the younger Vectis Form ...
of the United Kingdom. Furthermore, the prefrontal bone has a feature seen in ''Baryonyx'' but not allosauroids. Given the discrepancies with the placement of ''Eocarcharia'' as a carcharodontosaur and the many similarities to ''Suchomimus'', ''Ceratosuchops'', and ''Riparovenator'', the researchers considered it to be most parsimoniously regarded as a spinosaurid. Cau and Paterna's phylogenetic results placed ''Eocarcharia'' as the sister taxon to ''Suchomimus''. Both species differ in fourteen characters, precluding their taxonomic synonymization. This also provides further evidence to an observed trend of at least two spinosaurids coexisting in one ecosystem (e.g., ''Ceratosuchops'' and ''Riparovenator'' in the Wessex Formation). Since the maxilla OTU was recovered as distinct from the skull roof + holotype postorbital OTU, Cau and Paterna suggested that this bone—which demonstrates apparent allosauroid anatomy consistent with carcharodontosaurids—should be referred to a new taxon. Abbreviated results of their phylogenetic analysis are displayed in the cladogram below, with both ''Eocarchia'' OTUs indicated: the skull roof and holotype within the spinosaurid clade
Ceratosuchopsini Baryonychinae is an extinct clade or subfamily of spinosaurids from the Early Cretaceous of Europe and West Africa. The clade was named by Charig & Milner in 1986 and defined by Sereno et al. in 1998 and Holtz et al. in 2004 as all taxa more close ...
and the maxilla as a basal carcharodontosaurid. ⊞ buttons can be clicked to expand nodes. Notably, spinosaurid similarities have been noted prior to the Cau and Paterna's 2025 analysis; in 2022, Sereno and colleagues figured a newly-discovered skull roof of ''Suchomimus''. The similarity of this specimen to the postorbital and frontal of ''Eocarcharia'' was noted by Schade and colleagues the following year. In early 2025, Kellermann, Cuesta & Rauhut reinforced this comparison, claiming that the specimen may even be referrable to the genus. Cau and Paterna acknowledged these similarities, but noted characters more consistent with ''Suchomimus'' than ''Eocarcharia''. They also commented on the possibility that some of the other specimens traditionally referred to ''Suchomimus'' may actually belong to ''Eocarcharia''.


Paleobiology

Due to the fragmentary, chimeric, and indeterminate nature of the fossils of ''Eocarcharia'', little can be directly known about its
paleobiology Paleobiology (or palaeobiology) is an interdisciplinary field that combines the methods and findings found in both the earth sciences and the life sciences. An investigator in this field is known as a paleobiologist. Paleobiology is closely ...
. In their 2008 description, Sereno and Brusatte noted the thickness and robustness of the postorbital bosses. The brow was likely covered in
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
, extending its size. It also displays large, intricate sutures between the postorbital and frontal that suggest it would be stable against lateral impacts. The laterosphenoid's head is placed in a socket in the postorbital which, although shallower than that of advanced carcharodontosaurids, would be able to handle stress. This combination of stress adaptations illustrate that the brow was for more than purely display purposes. This led the authors to hypothesize that the postorbital brows of ''Eocarcharia'' and carcharodontosaurids were used for
intraspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organism ...
lateral
head-butting A headbutt or butt is a targeted strike with the head, typically involving the use of robust parts of the headbutter's cranium as the area of impact. The most effective headbutts strike the most sensitive areas of an opponent, such as the nose, ...
. This brow is absent in allosauroids and most spinosaurids, however the spinosaurids ''
Ceratosuchops ''Ceratosuchops'' (meaning "horned crocodile face") is a genus of spinosaurid from the Early Cretaceous (Barremian) of Britain. Discovery and naming The holotype remains of this taxon consist of IWCMS 2014.95.5 (premaxillary bodies), IWCM ...
'' and ''
Riparovenator ''Riparovenator'' ("riverbank hunter") is a genus of Baryonychinae, baryonychine spinosaurid dinosaur from the Early Cretaceous (Barremian) period of Britain. The genus contains a Monotypic taxon, single species, ''Riparovenator milnerae''. Di ...
'' also preserve large orbital bosses comparable to those in ''Eocarcharia''. Although present in large
tyrannosaurids Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera i ...
, the postorbital bosses of ''Eocarcharia'' and carcharodontosaurids project laterally, further suggesting that they played a role in lateral head-butting.


Paleoecology

''Eocarcharia'' is known from the Elrhaz Formation of the Tegama Group in an area of the Nigerian Ténéré Desert called Gadoufaoua. The Elrhaz Formation consists mainly of
fluvial A river is a natural stream of fresh water that flows on land or inside caves towards another body of water at a lower elevation, such as an ocean, lake, or another river. A river may run dry before reaching the end of its course if it ru ...
sandstones with low relief, much of which is obscured by sand dunes. The
sediments Sediment is a solid material that is transported to a new location where it is deposited. It occurs naturally and, through the processes of weathering and erosion, is broken down and subsequently sediment transport, transported by the action of ...
are coarse- to medium-grained, with almost no fine-grained horizons. ''Eocarcharia'' lived about 120 to 112 million years ago, during the
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
to
Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...
ages of the mid-Cretaceous. It likely lived in habitats dominated by inland
floodplains A floodplain or flood plain or bottomlands is an area of land adjacent to a river. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high discharge.Goudie, ...
(a
riparian zone A riparian zone or riparian area is the interface between land and a river or stream. In some regions, the terms riparian woodland, riparian forest, riparian buffer zone, riparian corridor, and riparian strip are used to characterize a ripari ...
). ''Eocarcharia'' lived alongside several other dinosaurs. These included other theropods, such as the spinosaurid and probable close relative ''Suchomimus'', ''Kryptops'' (known from a chimeric combination of
abelisaurid Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are foun ...
and
allosauroid 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 Carn ...
material), and the putative noasaurid ''
Afromimus ''Afromimus'' (meaning "Africa mimic") is a genus of ceratosaurian theropod dinosaur from the Early Cretaceous Elrhaz Formation of Niger. It contains a single species, ''A. tenerensis'', named in 2017 by Paul Sereno from parts of the right leg, ...
''. Several megaherbivores like the hadrosauriforms ''
Ouranosaurus ''Ouranosaurus'' is a genus of herbivorous basal hadrosauriform dinosaur that lived during the Aptian stage of the Early Cretaceous of modern-day Niger and Cameroon. ''Ouranosaurus'' measured about long and weighed . Two rather complete fossils ...
'' and ''
Lurdusaurus ''Lurdusaurus'' ("heavy lizard") is a genus of massive and unusually shaped iguanodont dinosaur from the Elrhaz Formation in Niger. It contains one species, ''L. arenatus''. The formation dates to the Early Cretaceous, roughly 112 million years ...
'',
dryosaurid Dryosauridae was a family of primitive iguanodonts, first proposed by Milner & Norman in 1984. They are known from Middle Jurassic to Early Cretaceous rocks of Africa, Europe, and North America. Dryosauridae was first proposed in 1984 by Britis ...
''
Elrhazosaurus ''Elrhazosaurus'' (meaning " Elrhaz lizard") is a genus of basal iguanodontian dinosaur, known from isolated bones found in Early Cretaceous rocks of Niger. These bones were initially thought to belong to a species of the related dryosaurid '' ...
'', and two
sauropods Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
, the
rebbachisaurid Rebbachisauridae is a Family (biology), family of sauropod dinosaurs known from fragmentary fossil remains from the Cretaceous of South America, Africa, North America, Europe and possibly Central Asia. Taxonomy In 1990 sauropod specialist Jack M ...
''
Nigersaurus ''Nigersaurus'' () is a genus of rebbachisaurid sauropod dinosaur that lived during the middle Cretaceous period, about 115 to 105 million years ago. It was discovered in the Elrhaz Formation in an area called Gadoufaoua, in Niger. Fossils o ...
'' and an unnamed
titanosaur Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still thr ...
, have been unearthed from Gadoufaoua. Together, these comprise one of the few associations of megaherbivores with a balance of sauropods and large
ornithopods Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivore, herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and rel ...
.
Crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. Extinct crocodylomorphs were considerably more ...
like ''
Sarcosuchus ''Sarcosuchus'' (), from Ancient Greek σάρξ (''sárx''), meaning "flesh", and σούχος (''soúkhos''), meaning "crocodile", is an extinct genus of crocodyliform a that lived during the Early Cretaceous, from the late Hauterivian to the ...
'', ''
Anatosuchus ''Anatosuchus'' ("duck crocodile", the name from the Latin ''anas'' ("duck") and the Greek ''souchos'' ("crocodile"), for the broad, duck-like snout) is an extinct genus of notosuchian crocodyliforms discovered in Gadoufaoua, Niger, and described ...
'', ''
Araripesuchus ''Araripesuchus'' is a genus of extinct crocodyliform that existed during the Cretaceous period of the late Mesozoic era some 125 to 66 million years ago. ''Araripesuchus'' is generally considered to be a notosuchian (belonging to the clade Mes ...
'', and ''
Stolokrosuchus ''Stolokrosuchus'' is an extinct genus of crocodyliforms found in the Early Cretaceous Elrhaz Formation, Niger. Discovery and naming The holotype includes a skull with a long thin snout and bony knobs on the prefrontal, and it was found in t ...
'' also lived there. In addition, remains of an
ornithocheirid Anhangueridae (alternatively called Ornithocheiridae, meaning "bird hands") is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the ...
pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
,
turtles Turtles are reptiles of the order Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked turtle ...
,
bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
, a
hybodont Hybodontiformes, commonly called hybodonts, are an extinct group of shark-like cartilaginous fish (chondrichthyans) which existed from the late Devonian to the Late Cretaceous. Hybodonts share a close common ancestry with modern sharks and rays ( ...
shark, and
bivalves Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consis ...
have been found. The aquatic fauna consists entirely of freshwater inhabitants.


See also

* ''
Kryptops ''Kryptops'' (meaning "covered face") is an extinct genus of abelisaurid theropod dinosaurs from the Early Cretaceous of Niger. It is known from a partial maxilla (upper jaw bone) found at the Gadoufaoua locality in the western Ténéré Desert, ...
'' *
2008 in paleontology Protozoa New taxa Plants Ferns and fern allies Angiosperms Monocots Arthropoda Arachnomorphs Crustaceans Research * Schram (2008) publishes an adjustment to the taxonomy of Paleozoic stomatopods, erecting the family Archaeocari ...


References

{{Taxonbar, from1=Q29574600, from2=Q132487 Carcharodontosauridae Dinosaur genera Albian dinosaurs Fossil taxa described in 2008 Taxa named by Paul Sereno Taxa named by Stephen L. Brusatte Dinosaurs of Niger