| Tameryraptor | |
|---|---|
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| Photograph of the holotype before its destruction in 1944 | |
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| Skeletal reconstruction of the holotype with known material in white | |
Scientific 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 anextinct genus of largecarcharodontosauriddinosaur that lived during theLate Cretaceous (Cenomanian age) in what is now Egypt. It is known from a partial skeleton collected in rock layers from theBahariya Formation by crews of German paleontologistErnst Stromer in 1914, comprising an incomplete skull, vertebrae, and several other postcranial elements. Stromer described the specimen in 1931, referring it to the previously namedCarcharodontosaurus on the basis of its tooth anatomy. In 1944, it was destroyed in theBombing of Munich during theSecond World War. The specimen remained assigned toCarcharodontosaurus saharicus until a review of photographs of the fossil material in 2025 allowed researchers to recognize the material as belonging to a distincttaxon known from asingle species,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 was distinct in that it bore a distinctive horn-like protuberance on its snout. Itsvertebrae were sturdy but contain depressions whereair sacs would be present. Several other gigantic theropods are known from Egypt during this period, including thespinosauridSpinosaurus, the controversialBahariasaurus, and unnamed largeabelisaurids.
In early April 1914, theropod fossils were found in marls nearAin Gedid, Egypt byAustro-Hungarian paleontologistRichard Markgraf. The sediments from this region derive from theCenomanian-agedBahariya Formation, one of many Cretaceous-aged sites ofNorth Africa.[1][2][3]: 162 Markgraf extensively collected dinosaur skeletons in Bahariya for his employer, German paleontologistErnst Stromer of thePaläontologisches Museum München (Bavarian State Collection of Paleontology). This Egyptian skeleton (specimen numberSNSB-BSPG 1922 X 46) consisted of several elements, most notably a partial skull which included the left and rightnasals, much of themaxillae (upper jaw bones), an incompletebraincase, andteeth. As for thepostcranial skeleton, threecervical (neck) vertebrae, onecaudal (tail) vertebra, twochevrons, a partialpelvis, the proximal portion of a dorsalrib, bothfemora (thigh bones), and the leftfibula (shin bone) were found.[1][4]
Due topolitical tensions between theGerman Empire and thenBritish-owned Egypt, this specimen took years to get to Germany. It was not until 1922 that the bones were transported toMunich, where Stromer described them in 1931. 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.[1]World War II broke out in 1939, leading to SNSB-BSPG 1922 X 46 and other Bahariya material to be destroyed during aBritish bombing raid on Munich during the night of April 24/25, 1944.[5][6] A braincaseendocast was made that survived the war, making it the only remaining relic of the specimen.[3]: 162, 164
The 1990s witnessed a resurgence in carcharodontosaurid research and discoveries, with the description of a novelCarcharodontosaurus saharicus specimen from Morocco in 1996.[2] This specimen was then designated theneotype of the species in 2007, though SNSB-BSPG 1922 X 46 was still classified as belonging to the species.[7] In 2025, Kellermann, Cuesta & RauhutdescribedTameryraptor 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. Thegeneric name,Tameryraptor, combinesTa-mery, an informalancient Egyptian name for the country meaning'beloved land', with theLatin wordraptor, meaning'thief'. Thespecific name,markgrafi, honors Richard Markgraf, the discoverer of the remains.[4]
In 1931, Stromer estimated that theTameryraptor holotype represented an individual similar in size to thetyrannosauridGorgosaurus,[1] which has been estimated at 8–9 metres (26–30 ft) in length.[8][9]
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 were 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 was accentuated by a depression behind the protrusion. While comparable to the nasal horn ofCeratosaurus, a distantly related theropod, it is much less pronounced. The nasals bear a large, extended antorbital fossa (depressions on the nasal) that extends along theantorbital fenestra (a large opening in the skull in front of the eye). This feature is one of several that distinguishesTameryraptor from other genera of carcharodontosaurid. Both maxillae were 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 ofC. saharicus is much larger. The maxillae'slateral (left and right) surfaces were adorned with rugosities, vertical ridges, and furrows that were much more pronounced than in related taxa. Based on the number ofalveoli (tooth sockets), the maxillae bore either 12 or 13 teeth,[4] a lower count than inCarcharodontosaurus.[2]
The rear portion of the skull was represented by theparietals (side and roof of cranium),frontals (front-top of cranium), part of thesupraoccipital (bottom rear of cranium), and partialocciput (region in the lower back portion of the cranium). Overall, the braincase andposterior (back portion) skull were comparable to those ofCarcharodontosaurus,Acrocanthosaurus,Giganotosaurus, andMeraxes. The frontals were 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 ofMeraxes. Theprefrontal facet (area where the frontal joins the prefrontal) on the frontal was not expanded, a characteristic unique toTameryraptor among Carcharodontosauridae. The maxillary teeth ofTameryraptor were more symmetrical and triangular than those ofCarcharodontosaurus, similar to a tooth fragment from theKem Kem Group in Morocco and an isolated tooth that has been assigned toMapusaurus. Like the teeth of other carcharodontosaurids, those ofTameryraptor bore horizontalenamel wrinkles.[4]
TheTameryraptor holotype was initially interpreted as one of the most complete postcranial specimens ofCarcharodontosaurus. This specimen preserved threecervical vertebrae, which were weathered severely. One was theaxis and the other two wereanterior (front) cervicals that were larger than the axis. The axis was wider than tall, though was incomplete, and bears parapophyses about halfway on its height on both of its sides. Its shortneural spine was inclined backwards and closely resembles the axes of derived carcharodontosaurids, but contrasts with those of the high-spinedNeovenator andAcrocanthosaurus. The cervical vertebrae, similar to the relatedGiganotosaurus, were topped by low neural spines joined with sturdytransverse processes which hung over thepleurocoels (shallow depressions on the sides ofcentra), which would contain pneumatic air sacs to lighten the vertebrae. The centra of these vertebrae have keels along their ventral sides. The middle cervical vertebra preserved, possibly the 4th position cervical, wasopisthocoelous (convex anterior ends of central, concave posterior ends) as in carcharodontosaurines. The ratio of transversal width to anterior centrum height was 1.3, making the vertebra much wider than the cervical vertebrae preserved in carcharodontosaurids likeGiganotosaurus andTyrannotitan but more similar to that ofAllosaurus, a morebasal theropod. The third cervical vertebra described was unfigured and poorly preserved, though was apparently more posterior along the neck. An anteriorcaudal vertebra was also known, which wasplatycoelous (flat anterior and posterior ends) and short. This vertebra was incomplete, missing much of the neural spine. The sides of its centrum were pleurocoelus as well. The caudal vertebra was strongly pneumatized, with the centrum covered in pneumaticforamina (openings in bone), a trait potentially diagnostic ofTameryraptor. Twohaemal arches, or chevrons, were preserved in this individual as well.[1][4]
The pelvis was incomplete, containing bothpubes and the leftischium. Uniquely, the ischium 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. The public shafts were strongly curved laterally, a condition observed in some related taxa, but from anterior view lacked intense curvature. As for the ischium, it was very incomplete, preserving primarily the proximal portion, but was firmly pointed downward. 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, though it was damaged during its transport to Germany. Thelesser trochanter (a projection from the shaft of the femur) well developed with a strong separation from the larger greater trochanter. Additionally, thefourth trochanter is similarly well developed and clearly visible in illustrations and photographs of the specimen. These traits are typical of basal carcharodontosaurids but contrast with the weakly developed lesser and fourth trochanters ofGiganotosaurus andMapusaurus. However, theaccessory trochanter is "spike-like", a trait distinguishingTameryraptor from other carcharodontosaurids. 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 bulgingcondyles whereas the posterior end was rounded, a trait distinguishingTameryraptor from other carcharodontosaurids.[1][4]
Tameryraptor is a genus in the family Carcharodontosauridae but, like the generaAcrocanthosaurus andLajasvenator, is classified outside of the subfamily Carcharodontosaurinae.[4] The fossils now assigned toTameryraptor were used by Stromer to create Carcharodontosauridae, a clade that originally only includedCarcharodontosaurus andBahariasaurus. Stromer noted the likeness ofTameryraptor, then assigned toCarcharodontosaurus, bones to the American theropodsAllosaurus andTyrannosaurus, leading him to consider the family part of Theropoda.[1] Carcharodontosauridae has since been expanded, with genera described from theJurassic and or Cretaceous of every continent exceptAntarctica.[10][2] The peak of diversity and size of the group was during the mid-Cretaceous, with members likeGiganotosaurus andCarcharodontosaurus itself reaching over 10 meters in length.[8]
In theirphylogenetic analyses, Kellermann, Cuesta & Rauhut (2025) recoveredTameryraptor as a non-carcharodontosaurine member of the Carcharodontosauridae in both analyses. This demonstrates thatTameryraptor was more basal than the highly derived carcharodontosaurids from South America likeMeraxes andTyrannotitan, but also more basal thanCarcharodontosaurus. Additionally, both phylogenetic analysis resulted inTameryraptor being at a similargrade asAcrocanthosaurus and other basal carcharodontosaurids likeLusovenator. The results of their analysis using merged OTUs (operational taxonomic units) is displayed in thecladogram below:[4]
| Carcharodontosauriformes |
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Gigantism in theropods evolved independently in separate groups (convergent evolution), withRodolfo Coria andLeonardo Salgado suggesting it was linked to common conditions in their environments orecosystems.[11] 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 theEarly Cretaceousperiod.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.[12][2] Previously, it was thought that the Cretaceous world wasbiogeographically separated, with the northern continents being dominated by tyrannosaurids, South America by abelisaurids, and Africa by carcharodontosaurids.[13][14][15]
The description ofTameryraptor 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.[16][17][18] 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 othertetrapods from the two sites but the supposed overlap of genera likeSpinosaurus andCarcharodontosaurus 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]
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The function of the small nasal horn ofTameryraptor was discussed in its description, with the possibility of its presence being a result of sexual dimorphism. However, this was dismissed as no other theropod genera with crests or horns, such asCeratosaurus, Allosaurus, andMajungasaurus, preserve definitive sexual dimorphism in their skull ornamentation.[4] Other theropods such asCarcharodontosaurus, Allosaurus, andAcrocanthosaurus too have enlarged crests, whose purpose is unknown. Paleontologist Daniel Chure hypothesized that these crests were used for "head-butting" between individuals, but how durable they are has not been studied.[19]
The dentition of allosauroids is distinct, with carcharodontosaurid teeth bearing distinctly thin and blade-like teeth. However, these teeth are thin and likely could not sustain impact against hard surfaces like bone without potentially bending and snapping. This danger is exacerbated by the straight edges, slightly recurved tips, andsinusoidal shapes observed in their dentition. Despite these traits, the teeth are still much more robust than those of smaller theropods and due to their overall size could take more pressure.[20][21][22]
North Africa during the Cenomanian stage of the Late Cretaceous bordered theTethys Sea, which transformed the region into amangrove-dominated coastal environment filled with vasttidal flats andwaterways.[23]Tameryraptor is known from theBahariya Formation, then awetland environment, alongside the coevalSpinosaurus 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 ofTameryraptor suggested that such superficial comparisons require further examination.[4] Contemporaryabelisaurid dinosaurs from the Bahariya Formation were also terrestrial carnivores, preying on other terrestrial fauna.[24] Some sauropods are also known from the same formation such asParalititan andAegyptosaurus.[25] 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 thepleurodianApertotemporalis, large bony fish likeMawsonia[26] andParanogmius,[27]sawskatesOnchopristis andSchizorhiza,[28]sharks likeSqualicorax andCretolamna, and a broad selection ofinvertebrates.[29]
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.[30] In North Africa, carcharodontosaurids are represented byC. saharicus andSauroniops in the Kem Kem Beds,Eocarcharia and potentiallyCarcharodontosaurus in theElrhaz Formation, andC. iguidensis in theEchkar Formation.[7][4]
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