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Eocarcharia

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Eocarcharia
Temporal range: Early Cretaceous
(AptianAlbian), ~112 Ma
Holotype postorbital
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Family: Spinosauridae (?)
Subfamily: Baryonychinae (?)
Genus: Eocarcharia
Sereno & Brusatte, 2008
Type species
Eocarcharia dinops
Sereno and Brusatte, 2008

Eocarcharia (meaning "dawn shark") is an extinct genus of theropod dinosaurs known from what is now Niger. It is known from several postorbitals (bones behind the orbit) and an incomplete skull roof found at the Gadoufaoua locality in the western Ténéré Desert, in rocks of the Early Cretaceous (AptianAlbian ages) Elrhaz Formation. The fossils were collected in 2000 by an expedition led by American paleontologist Paul Sereno. They were then described in 2008 by Sereno and Steve Brusatte. The genus contains a single species, Eocarcharia dinops. Sereno and Brusatte also referred a maxilla (upper jaw bone) and teeth to this species, but later studies suggested that it is chimaeric, comprising bones of multiple unrelated taxa. Some of the Eocarcharia material, including the holotype specimen, likely belongs to a baryonychine spinosaurid. This would render Eocarcharia a member of this group, closely related to the coeval Suchomimus, while the definitively carcharodontosaurid maxilla and teeth belong to a separate distinct taxon.

Little is known about Eocarcharia due to its fragmentary and chimeric nature. When referred to Carcharodontosauridae, it was estimated to be 6–8 metres (20–26 ft) long, making it smaller than derived carcharodontosaurids like Giganotosaurus, and Carcharodontosaurus. The postorbital is robust and bears a large brow, a distinct characteristic of the dinosaur. It was possibly covered in keratin and used for lateral head-butting with other individuals of its species. Eocarcharia lived in a fluvial environment with vast floodplains alongside many other dinosaurs, pterosaurs, crocodylomorphs, and freshwater animals.

Discovery and naming

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Location of Gadoufaoua in Niger
Reconstruction of the likely chimaeric hypodigm skull of Eocarcharia based on the initial interpretation of the material

In 2000, American paleontologist Paul Sereno led an expedition conducted with the University of Chicago and funded by the National Geographic Society. The purpose of this trip was to explore fossiliferous sandstone outcrops in a site on the western edge of the Nigerian Ténéré Desert known as Gadoufaoua.[1][2] These layers belong to the Elrhaz Formation, to the Early Cretaceous. During the expedition, several isolated theropod skull bones were collected, including a maxilla (main tooth-bearing bone of the upper jaw) and two maxilla fragments, five postorbitals (posterior orbit bones), a left frontal and prefrontal (skull roof bones), and five teeth. 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.[2]

Sereno believed these remains all belonged to the same species, and in a 2004 paper coauthored with Jeffrey Wilson and Jack Conrad, mentioned them as belonging to an undescribed carcharodontosaurid.[3] Four years later, Sereno and Steve Brusatte 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 (name-bearing) specimen. The generic name derives from the Ancient Greek words eos, meaning 'dawn', and karcharias, meaning 'shark', referencing the early-diverging nature of Eocarcharia in relation to its 'shark-toothed' relatives. The specific name, dinops, is a Greek term meaning "fierce-eyed", referring to the postorbital ornamentation above the eye.[2][4]

Description

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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 confidently known remains of Eocarcharia consist of the holotype postorbital, four referred postorbitals, a left frontal, and a left prefrontal.[5] Referencing the more complete skulls and skeletons of other carcharodontosaurids like Acrocanthosaurus and Carcharodontosaurus, they determined that Eocarcharia was likely about 6–8 metres (20–26 ft) long, half the linear size of the largest, more derived members of the family.[2]

Posterior skull

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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, a trait noted by Sereno and Brusatte to be diagnostic (a distinct feature of a species). However, the occurrence of an exaggerated orbital boss is also found in spinosaurids.[6][5] An orbital boss is also found in Mapusaurus and Giganotosaurus, though the orbital bosses of these genera are ossified 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.[2][7]

Skull roof referred to Eocarcharia in dorsal and ventral (left) and lateral and medial (right) view

The margins of the postorbital make up segments of the orbit edges, the laterotemporal fenestra, and the supratemporal fenestra. Several of the postorbitals known from Eocarcharia preserve their lacrimal articular surfaces (part of the postorbital that articulates with the lacrimal), which are 9 millimetres (0.35 in) deep and 12 millimetres (0.47 in) 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 (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 jugal (cheekbone), whereas carcharodontosaurids like Carcharodontosaurus have broader facets. Midshafts of the ventral (bottom) ramus (a branch of bone) in spinosaurids and megalosaurids are typically U-shaped in cross-section, 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.[2][5]

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 fossa (a depression along the temporal fenestra) is greatly exposed, another diagnostic characteristic. In ventral view, the anterior portion of the frontal is exposed and forms the roof of the olfactory section of the endocranium (part of skull that contains the brain). This portion of the frontal is narrow, in contrast to those of tetanurans 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,[8] whereas in genera like Acrocanthosaurus, Eocarcharia, and Concavenator, they are unfused.[7] 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.[2]

Classification

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As a carcharodontosaurid

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In their phylogenetic analyses, 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 to the North American Acrocanthosaurus.[2]

Carcharodontosaurid maxilla referred to Eocarcharia

In the same 2008 paper, Sereno and Brusatte named Kryptops palaios on the basis of an abelisaurid 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.[2] In 2012, Matthew Carrano and colleagues considered Kryptops palaios to be a chimera (specimen composed of multiple species), and stated that its postcranial remains, especially the pelvis and sacrum, may actually belong to a carcharodontosaurid, possibly Eocarcharia.[9] However, these bones do not overlap with the Eocarcharia holotype, which consists only of an isolated postorbital.[2] This hypothesis was supported by later studies, who agreed that the postcranial remains belonged to an allosauroid,[10] carcharodontosaurid,[11][12] or a metriacanthosaurid.[5]

In their 2024 description of a new specimen of the giant carcharodontosaurid Taurovenator, Rolando et al. recovered Eocarcharia as an early-diverging member of the clade, branching after Neovenator. The results of their phylogenetic analyses are displayed in the cladogram below:[13]

Carcharodontosauridae

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, 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 units (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. 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.[14]

As a spinosaurid

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Reconstructed skull of the spinosaurid Suchomimus, a possible close relative of Eocarcharia

In 2024, Andrea Cau published the results of a comprehensive theropod phylogenetic framework.[10] 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 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 (unique derived traits) proposed for Eocarcharia by Sereno and Brusatte are also uniquely shared with Ceratosuchops, a baryonychine from the Wessex Formation of the United Kingdom.[6] 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.[5]

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 and the maxilla as a basal carcharodontosaurid. ⊞ buttons can be clicked to expand nodes.[5]

Carnosauria
Megalosauroidea
phylogenetic position of Eocarcharia (holotype + skull roof)
Allosauroidea
phylogenetic position of the referred "Eocarcharia" maxilla

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.[15] The similarity of this specimen to the postorbital and frontal of Eocarcharia was noted by Schade and colleagues the following year.[16] In early 2025, Kellermann, Cuesta & Rauhut reinforced this comparison, claiming that the specimen may even be referrable to the genus.[14] Cau and Paterna acknowledged these similarities, but noted characters more consistent with Suchomimus than Eocarcharia.[5]

Paleobiology

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Due to the fragmentary, chimeric, and indeterminate nature of the fossils of Eocarcharia, little can be directly known about its paleobiology. In their 2008 description, Sereno and Brusatte noted the thickness and robustness of the postorbital bosses. The brow was likely covered in keratin, 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 lateral head-butting. This brow is absent in allosauroids and most spinosaurids, however the spinosaurids Ceratosuchops and Riparovenator also preserve large orbital bosses comparable to those in Eocarcharia.[6][5] Although present in large tyrannosaurids, the postorbital bosses of Eocarcharia and carcharodontosaurids project laterally,[14][11] further suggesting that they played a role in lateral head-butting.[2]

Paleoecology

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Restoration of Suchomimus (foreground) and Nigersaurus (background) in the Elrhaz Formation environment

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 sandstones with low relief, much of which is obscured by sand dunes. The sediments are coarse- to medium-grained, with almost no fine-grained horizons.[17] Eocarcharia lived about 120 to 112 million years ago, during the Aptian to Albian ages of the mid-Cretaceous.[2] It likely lived in habitats dominated by inland floodplains (a riparian zone).[18]

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 and allosauroid material),[5] and the putative noasaurid Afromimus.[19] Several megaherbivores like the hadrosauriforms Ouranosaurus and Lurdusaurus, dryosaurid Elrhazosaurus, and two sauropods, the rebbachisaurid Nigersaurus and an unnamed titanosaur, have been unearthed from Gadoufaoua. Together, these comprise one of the few associations of megaherbivores with a balance of sauropods and large ornithopods. Crocodylomorphs like Sarcosuchus, Anatosuchus, Araripesuchus, and Stolokrosuchus also lived there. In addition, remains of an ornithocheirid pterosaur, turtles, bony fish, a hybodont shark, and bivalves have been found. The aquatic fauna consists entirely of freshwater inhabitants.[18][20][21]

References

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  1. ^ Koppes, Steve (2000). "Dinosaur Expedition 2000 allows us to join team in Niger". chronicle.uchicago.edu. Retrieved 15 May 2025.
  2. ^ a b c d e f g h i j k l Sereno, Paul C. and Brusatte, Stephen L. (January 2008). "Basal Abelisaurid and Carcharodontosaurid Theropods from the Lower Cretaceous Elrhaz Formation of Niger". Acta Palaeontologica Polonica. 53 (1): 15–46. doi:10.4202/app.2008.0102. hdl:20.500.11820/5d55ed2e-52f2-4e4a-9ca1-fd1732f2f964. ISSN 0567-7920.
  3. ^ Sereno, Paul C.; Wilson, Jeffrey A.; Conrad, Jack L. (7 July 2004). "New dinosaurs link southern landmasses in the Mid–Cretaceous". Proceedings of the Royal Society of London. Series B: Biological Sciences. 271 (1546): 1325–1330. doi:10.1098/rspb.2004.2692. PMC 1691741. PMID 15306329.
  4. ^ "Discoveries | Paul Sereno - Paleontologist | The University of Chicago". paulsereno.uchicago.edu. Retrieved 15 May 2025.
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  6. ^ a b c Barker, C.T.; Hone, D.; Naish, D.; Cau, A.; Lockwood, J.; Foster, B.; Clarkin, C.; Schneider, P.; Gostling, N. (2021). "New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae". Scientific Reports. 11 (1): 19340. Bibcode:2021NatSR..1119340B. doi:10.1038/s41598-021-97870-8. PMC 8481559. PMID 34588472.
  7. ^ a b Cuesta, Elena; Vidal, Daniel; Ortega, Francisco; Sanz, José L. (1 November 2018). "The cranial osteology of Concavenator corcovatus (Theropoda; Carcharodontosauria) from the Lower Cretaceous of Spain". Cretaceous Research. 91: 176–194. Bibcode:2018CrRes..91..176C. doi:10.1016/j.cretres.2018.06.007. ISSN 0195-6671.
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  14. ^ a b c Kellermann, Maximilian; Cuesta, Elena; Rauhut, Oliver W. M. (14 January 2025). "Re-evaluation of the Bahariya Formation carcharodontosaurid (Dinosauria: Theropoda) and its implications for allosauroid phylogeny". PLOS One. 20 (1): e0311096. Bibcode:2025PLoSO..2011096K. doi:10.1371/journal.pone.0311096. ISSN 1932-6203. PMC 11731741. PMID 39808629.
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  16. ^ Schade, Marco; Rauhut, Oliver; Foth, Christian; Moleman, Olof; Evers, Serjoscha (2023). "A reappraisal of the cranial and mandibular osteology of the spinosaurid Irritator challengeri (Dinosauria: Theropoda)". Palaeontologia Electronica. doi:10.26879/1242. S2CID 258649428.
  17. ^ Sereno, Paul C.; Beck, Allison L.; Dutheil, Didier B.; Larsson, Hans C. E.; Lyon, Gabrielle H.; Moussa, Bourahima; Sadleir, Rudyard W.; Sidor, Christian A.; Varricchio, David J.; Wilson, Gregory P.; Wilson, Jeffrey A. (12 November 1999). "Cretaceous Sauropods from the Sahara and the Uneven Rate of Skeletal Evolution Among Dinosaurs". Science. 286 (5443): 1342–1347. doi:10.1126/science.286.5443.1342. PMID 10558986.
  18. ^ a b Sereno, Paul C.; Wilson, Jeffrey A.; Witmer, Lawrence M.; Whitlock, John A.; Maga, Abdoulaye; Ide, Oumarou; Rowe, Timothy A. (21 November 2007). "Structural Extremes in a Cretaceous Dinosaur". PLOS ONE. 2 (11): e1230. Bibcode:2007PLoSO...2.1230S. doi:10.1371/journal.pone.0001230. ISSN 1932-6203. PMC 2077925. PMID 18030355.
  19. ^ Cerroni, Mauricio A.; Agnolin, Federico L.; Brissón Egli, Federico; Novas, Fernando E. (1 November 2019). "The phylogenetic position of Afromimus tenerensis Sereno, 2017 and its paleobiogeographical implications". Journal of African Earth Sciences. 159: 103572. Bibcode:2019JAfES.15903572C. doi:10.1016/j.jafrearsci.2019.103572. ISSN 1464-343X.
  20. ^ Sereno, Paul C. (November 2017). "Early Cretaceous Ornithomimosaurs (Dinosauria: Coelurosauria) from Africa". Ameghiniana. 54 (5): 576–616. Bibcode:2017Amegh..54..576S. doi:10.5710/AMGH.23.10.2017.3155. ISSN 0002-7014.
  21. ^ Sereno, Paul; Larsson, Hans (19 November 2009). "Cretaceous Crocodyliforms from the Sahara". ZooKeys (28): 1–143. Bibcode:2009ZooK...28....1S. doi:10.3897/zookeys.28.325. ISSN 1313-2970.