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Tameryraptor

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Tameryraptor
Temporal range: Late Cretaceous, Cenomanian
Photograph of the holotype before its destruction in 1944
Skeletal reconstruction of the holotype with known material in white
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: †Carcharodontosauria
Family: †Carcharodontosauridae
Genus: †Tameryraptor
Kellermann, Cuesta & Rauhut, 2025
Species:
†T. markgrafi
Binomial name
†Tameryraptor markgrafi
Kellermann, Cuesta & Rauhut, 2025

Tameryraptor ("thief from the beloved land") is an extinct genus of large carcharodontosaurid theropod dinosaur that lived from around 100 million to 94 million years ago during the Late Cretaceous (Cenomanian age) in the Bahariya Formation of Egypt. A partial skeleton containing an incomplete skull, vertebrae, and several other postrcranial elements was collected by crews of German paleontologist Ernst Stromer in 1914. Stromer described the specimen in 1931, referring them to the previously named Carcharodontosaurus on the basis of its teeth anatomy. In 1944, it was destroyed during the Bombing of Munich during the Second World War. The specimen remained assigned to Carcharodontosaurus saharicus until a subsequent review of photographs of the fossil material allowed researchers in 2025 to recognize the material as belonging to a distinct taxon, naming it Tameryraptor markgrafi.

Tameryraptor is one of many large carcharodontosaurid dinosaurs. It was one of the only African carcharodontosaurids found that preserved associated cranial and postcranial remains Like its relatives, it had a large, lightly-built skull, but is distinct in that it bore a distinctive horn-like protuberance on its snout. Its vertebrae are sturdy but contain depressions where air sacs systems would be present. Many other gigantic theropods are known from Egypt during this period, including the spinosaurid Spinosaurus, the controversial Bahariasaurus, and unnamed large abelisaurids. Egypt at the time was blanketed in mangrove forests and wetlands, creating a hotspot of fish, crocodyliform, and pterosaur diversity.

Discovery and naming

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Stromer's (now outdated) reconstruction of the skull of Carcharodontosaurus saharicus using the Tameryraptor holotype

In early April 1914, theropod fossils were found in marls near Ain Gedid, Egypt by Austro-Hungarian paleontologist Richard Markgraf. The sediments from this region derive from the Cenomanian-aged Bahariya Formation, one of many Cretaceous-aged sites of North Africa.[1][2][3] Markgraf extensively collected dinosaur skeletons in Bahariya for his employer, German paleontologist Ernst Stromer of the Paläontologisches Museum München (Bavarian State Collection of Paleontology). This Egyptian skeleton (SNSB-BSPG 1922 X 46) consisted of several elements, most notably a partial skull. This skull material included: the left and right nasals, much of the maxillae, an incomplete braincase, and teeth. As for the postcranial skeleton, three cervical vertebrae and a caudal vertebra, two chevrons, a partial pelvis, the proximal portion of a dorsal rib, both femora, and the left fibula.[3][4]

Due to political tensions between the German Empire and then British-owned Egypt, this specimen took years to get to Germany. It was not until 1922 that the bones were transported to Munich, where Stromer described them in 1931.[3] Stromer recognized that the teeth of this specimen matched the characteristic dentition of those described by Depéret and Savornin in 1925 for their new species "Megalosaurus" saharicus. He found it necessary to erect a new genus for this species, Carcharodontosaurus. World War II broke out in 1939, leading to SNSB-BSPG 1922 X 46 and other Bahariya material to be destroyed during a British bombing raid on Munich during the night of April 24/25, 1944.[5][6] A braincase endocast was made that survived the war, making it the only remaining relic of the specimen.[2]

The 1990s witnessed a resurgence in carcharodontosaurid research and discoveries, with the description of a novel Carcharodontosaurus saharicus specimen from Morocco in 1996. This specimen was then designated the neotype of the species in 2007, though SNSB-BSPG 1922 X 46 was still classified as belonging to the species.[1][7] In 2025, Kellermann, Cuesta & Rauhut described Tameryraptor markgrafi as a new genus and species of carcharodontosaurid theropods based on these fossil remains. Since the fossil remains were destroyed, they established their description based on the remaining endocast, archival photographs, and Stromer's initial drawings of the fossil material. The generic name, Tameryraptor, combines Ta-mery, an informal ancient Egyptian name for the country—meaning "beloved land"—with the Latin word "raptor", meaning "thief". The specific name, markgrafi, honors Richard Markgraf, the discoverer of the remains.[4]

Description

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Size of the Carcharodontosaurus neotype (dark yellow) and Tameryraptor holotype (bright yellow) compared to a human

In 1931, Stromer estimated that the Tameryraptor holotype represented an individual similar in size to the tyrannosaurid Gorgosaurus,[3] which has been estimated at 8–9 metres (26–30 ft) in length.[8][9]

Skull and dentition

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Speculative life restoration of Tameryraptor with a nasal horn covered in extensive keratin

The skull Stromer described was incomplete and severely damaged, with the snout represented only by the nearly-complete left and right nasals and the damaged left maxilla. The middle parts of the nasals bear strong rugosities, similar to those of other carcharodontosaurids. However, they are characterized by a horn-like protrusion, measuring 3 cm (1.2 in) in height, which is not observed in any other taxon. The horn's prominence is accentuated by a depression behind the protrusion. While comparable to the nasal horn of Ceratosaurus, a distantly related theropod, it is much less pronounced. The nasals bear a large, extended antorbital fossa that extends along the later surfaces of the bones. Both of the maxillae are preserved though feature damage and wear. The maxilla of SNSB-BSPG 1922 X 46 would have been 70 centimetres (28 in) long when complete, whereas that of C. saharicus is much larger. The maxillae's lateral surfaces are adorned with rugosities, vertical ridges, and furrows that are much more pronounced than in related taxa. Based on the number of alveoli (tooth socket), the maxillae bore either 12 or 13 teeth, a tooth count lower than that of Carcharodontosaurus.[1][4]

The rear portion of the skull of Tameryraptor is represented by the parietals (side and roof of cranium), frontals (front-top of cranium), part of the supraoccipital (bottom rear of cranium), and partial occipitals (bones relating to the ear). Overall, the braincase and posterior skull is comparable to those of Carcharodontosaurus, Acrocanthosaurus, Giganotosaurus, and Meraxes. The frontals are convex on their exposed portion in contrast to the flat frontals of related genera. The supraoccipital features a prominent dorsal crest that extends towards the paroccipital (lateral side of occipital) process, similar to that of Meraxes. The maxillary teeth of Tameryraptor are more symmetrical and triangular than those of Carcharodontosaurus, similar to a tooth fragment from the Kem Kem Group and an isolated tooth that has been assigned to Mapusaurus. Like the teeth of other carcharodontosaurids, those of Tameryraptor bore horizontal enamel wrinkles.[4]

Postcranial skeleton

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Stromer's reconstruction of the pelvis and left hindlimb of the Tameryraptor holotype

The Tameryraptor holotype was initially interpreted as one of the most complete postcranial specimens of Carcharodontosaurus. This specimen preserved three cervical vertebrae, which were weathered severely. One is the axis and the other two are articulated anterior cervicals that are larger than the axis. The cervical vertebrae, similar to the related Giganotosaurus, are topped by low neural spines joined with sturdy transverse processes which hung over the pleurocoels (shallow depressions on the sides of centra), which would contain pneumatic air sacs to lighten the vertebrae. The centra of these vertebrae are adorned by keels along their ventral sides. An anterior caudal vertebra was also known, which was platycoelous (flat anterior and posterior ends) and short. This caudal was incomplete, missing much of the neural spine. The sides of its centrum were pleurocoelus as well. A haemal arch was preserved in this individual as well.[3][4]

The pelvis was incomplete, containing both pubes and the left ischium. The ischium is uniquely pointed almost directly horizontally. The pubes were likely nearly 1 metre (3.3 ft) when fully preserved, with thin shafts that were transversely expanded at the anterior ends where they connected, creating a V-shape in anterior view. Both femora in addition to the left fibula were recovered, the former element being one of the largest recorded from a theropod at 1.26 metres (4.1 ft) in length. Its femora lacked strong curvature. The greater trochanter is small but has a notable protrusion, which would attach to the m. caudofemoralis longus muscle of the tail. Its fibula was only 88 centimetres (35 in) long, around 1/3rd the length of the femora. The anterior end was triangular in lateral view with bulging condyles whereas the posterior end is rounded.[3][4]

Classification

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Tameryraptor is a genus in the family Carcharodontosauridae but, like the genera Acrocanthosaurus and Lajasvenator, is classified outside of the subfamily Carcharodontosaurinae.[4] The fossils now assigned to Tameryraptor were used by Stromer to create Carcharodontosauridae, a clade that originally only included Carcharodontosaurus and Bahariasaurus. Stromer noted the likeness of Tameryraptor, then assigned to Carcharodontosaurus, bones to the American theropods Allosaurus and Tyrannosaurus, leading him to consider the family part of Theropoda.[3]

In their phylogenetic analyses, Kellermann, Cuesta & Rauhut (2025) recovered Tameryraptor as a non-carcharodontosaurine member of the Carcharodontosauridae. Their analyses found support for a sister taxon relationship of carcharodontosaurids and metriacanthosaurids, which the authors named as a new clade, Carcharodontosauriformes. The results of their analysis using merged OTUs (operational taxonomic units) is displayed in the cladogram below:[4]

Carcharodontosauriformes

Evolution

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Life reconstruction of Tameryraptor.
Speculative life restoration of Tameryraptor in its environment

Rodolfo Coria and Leonardo Salgado suggested that the convergent evolution of gigantism in theropods could have been linked to common conditions in their environments or ecosystems.[10] Sereno and colleagues found that the presence of carcharodontosaurids in Africa (Carcharodontosaurus), North America (Acrocanthosaurus), and South America (Giganotosaurus), showed the group had a transcontinental distribution by the Early Cretaceous period. Dispersal routes between the northern and southern continents appear to have been severed by ocean barriers in the Late Cretaceous, which led to more distinct, provincial faunas, by preventing exchange.[11][1] Previously, it was thought that the Cretaceous world was biogeographically separated, with the northern continents being dominated by tyrannosaurids, South America by abelisaurids, and Africa by carcharodontosaurids.[12]

The description of Tameryraptor highlighted the underestimated diversity of theropod taxa in North Africa during the mid-Cretaceous, with Kellermann, Cuesta & Rauhut stating that the Bahariya and Kem Kem theropods may not be conspecific but instead belong to distinct genera,[4] a view held in some prior studies.[13][14][15] Although the similarity of the ages of the two locales has been used to justify this view, in actuality little direct analysis or comparison of fossil material has been made. This is further complicated by the destruction of the Munich collection and the lack of in-depth descriptions of Moroccan material. The lack of genus overlap in other tetrapods from the two sites but the supposed overlap of genera like Spinosaurus and Carcharodontosaurus was also noted by the study. It is for these reasons that the researchers stated that the evolution of large theropod dinosaurs in similarily-aged strata in Brazil, Egypt, Morocco, and Argentina resulted in greater biodiversity of genera and species than previously hypothesized.[4]

Paleoecology

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Restoration of the fauna of the Bahariya Formation, with Tameryraptor in the center background

North Africa during the Cenomanian stage of the Late Cretaceous bordered the Tethys Sea, which transformed the region into a mangrove-dominated coastal environment filled with vast tidal flats and waterways.[16] Tameryraptor lived in the Bahariya Formation, then a wetland environment, alongside the coeval Spinosaurus which is also known from the Kem Kem beds. The faunal composition of both the Bahariya Formation and the Kem Kem beds were thought to be similar in the past, but the describers of Tameryraptor suggested that such superficial comparisons require further examination.[4] Contemporary abelisaurid dinosaurs from the Bahariya Formation were also terrestrial carnivores, preying on other terrestrial fauna.[17] Some sauropods are also known from the same formation such as Paralititan and Aegyptosaurus.[18] A diverse fauna of aquatic animals is known from the Bahariya Formation. Underwater life diversity exploded during this period in the mangroves of North Africa, with turtles represented by the pleurodian Apertotemporalis, large bony fish like Mawsonia[19] and Paranogmius,[20] sawskates Onchopristis and Schizorhiza,[21] sharks like Squalicorax and Cretolamna, and a broad selection of invertebrates.[22]

The composition of the dinosaur fauna of these sites is an anomaly, as there are fewer herbivorous dinosaur species relative to carnivorous dinosaurs than usual. This indicates that there was niche partitioning between the different theropod clades, with spinosaurids consuming fish while other groups hunted herbivorous dinosaurs.[23] It coexisted with the sauropods Paralititan and Aegyptosaurus which are found in the Bahariya Formation.[18] In North Africa, carcharodontosaurids are represented by C. saharicus and Sauroniops in the Kem Kem Beds, Eocarcharia and potentially Carcharodontosaurus in the Elrhaz Formation, and C. iguidensis in the Echkar Formation.[7][4]

References

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