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Deltadromeus

Deltadromeus is an extinct genus of controversial theropod dinosaur that lived in present-day Morocco during the mid-Cretaceous period. It was described by American paleontologist Paul Sereno and colleagues in 1996. The genus contains a single species, D. agilis, named based on an incomplete postcranial skeleton, the holotype specimen. However, some fossils from the Bahariya Formation of Egypt that were formerly referred to the theropod Bahariasaurus have been suggested to belong to Deltadromeus. The holotype specimen of Deltadromeus was unearthed by a joint expedition by the University of Chicago and the Service Géologique du Maroc in Errachidia Province, eastern Morocco in rock layers coming from the Gara Sbaa Formation of the Kem Kem Beds. This indicates that these fossils date to the Cenomanian stage of the Cretaceous period, 100-95 million years ago.

Discovery and naming
region and outcrops Fossils definitively assigned to Deltadromeus were first discovered in 1996 by American paleontologist Paul Sereno, who was part of a joint expedition by the University of Chicago and the Service Géologique du Maroc in an outcrop of the Kem Kem Beds known as Aferdou N'Chaft in Errachidia Province, eastern Morocco. This expedition was one of many that explored dinosaur-bearing outcrops in North Africa throughout the 1990s and 2000s. The remains found consisted of an incomplete postcranial skeleton, cataloged as UCRC PV11 at the University of Chicago before being transferred to the Ministry of Energy, Mines and Environment in Rabat and catalogued as SGM-Din 2, that had been found in a single site. The strata of this site is made up of sandstones which belong to the Gara Sbaa Formation of the Kem Kem Beds, which dates to the Cenomanian-Albian ages of the middle Cretaceous period. These expeditions recovered many theropod fossils, including an individual found in 1922 known from several vertebrae, a rib fragment, and an incomplete pelvis that German paleontologist Ernst Stromer described as belonging to a new genus and species of theropod, Bahariasaurus ingens, in 1934. The proximal right tibia had also been assigned to the allosauroid Erectopus, Sereno and colleagues maintained that Bahariasaurus and Deltadromeus were different genera, but the validity and synonymy of the two genera is controversial. In 2024, Belgian paleontologist Christophe Hendrickx and colleagues described several theropod teeth from the Kem Kem Beds, one of which they stated is from either a non-abelisauroid ceratosaur or a megaraptoran, possibly Deltadromeus itself. However, the lack of overlap and the controversy surrounding Deltadromeus affinities made this impossible to prove until more fossils are found. ==Description==
Description
compared to a human The holotype individual was estimated to have measured in length and weighed around , slightly more than an imperial ton,by Australian researcher Frank Seebacher in 2001. Assigned specimens from the Bahariya Formation come from a much larger individual, with the femur IPHG 1912 VIII 69 having a length of , compared to the femur of the holotype. These referred specimens, if the referral to Deltadromeus is validated, would indicate that members of the genus could grow up to in length, approximately the length of a Tyrannosaurus rex. Anatomy and forelimb|leftDeltadromeus is very poorly known, only being represented by an incomplete postcranial skeleton which is missing much of the vertebrae, ribs, and forelimbs. In addition to its complicated taxonomy, this makes the anatomy of Deltadromeus poorly understood. The tibia/femur length/diameter ratio for example is 0.95 in Deltadromeus, whereas it is only 0.76 in Allosaurus and 1.16 in Ornithomimus. This indicates that the hindlimbs are relatively long for a theropod. Additionally, the humerus length/femur length ratio is higher in Deltadromeus than genera like Allosaurus, indicating that the forelimbs are not as reduced in that genus and Tyrannosaurus. The coracoid and acromion processes are broadly expanded anteroposteriorly (front-back), a characteristic believed to be diagnostic of Deltadromeus ==Classification==
Classification
The phylogenetic position of Deltadromeus is complicated by the fragmentary nature of the holotype, though others have found it to be more primitive, possibly related to the more ancestral ceratosaurs Elaphrosaurus and Limusaurus. A more comprehensive study of noasaurid relationships published in 2016 effectively agreed with both of these interpretations, with Deltadromeus, Limusaurus and Elaphrosaurus all found to be within the Noasauridae. A 2017 paper describing ontogenetic changes in Limusaurus and the effect of juvenile taxa on phylogenetic analyses placed Deltadromeus as a noasaurid in every analysis regardless of which Limusaurus specimen was used, although the analyses did not include Gualicho or Aoniraptor. According to the authors, resolving the phylogenetic positions of Gualicho, Aoniraptor, Deltadromeus and megaraptorans is a critical issue facing theropod systematics. Deltadromeus was also considered a noasaurid in a 2020 review of the Kem Kem Group geology and fauna. Since then, some studies find Deltadromeus as a ceratosaur, The cladogram below follows the 2016 Gualicho analysis by Sebastián Apesteguía, Nathan D. Smith, Rubén Juarez Valieri and Peter J. Makovicky. In 2020, Ibrahim and colleagues acknowledged similarities between the two genera, but considered it unlikely that Deltadromeus represents a specimen of Bahariasaurus due to perceived differences in the pelvic bones. They further regarded Bahariasaurus as a nomen dubium. The following year, Cau and Paterna used an updated version of this dataset to reanalyze the relationships of Bahariasaurus, Deltadromeus, and other Cretaceous theropods from Africa. They determined that the variation observed between specimens of Deltadromeus and Bahariasaurus was the result of individual and ontogenetic variation, as the former is known from immature remains. They further reidentified specimen SNSB-BSPG1912VIII82—recognized as a indeterminate theropod pubis by Stromer in his 1934 description of Bahariasaurus—as a complete ischium. The authors observed anatomical characters that the bone shares with the less complete ischia of the holotypes of both Bahariasaurus and Deltadromeus, which they used to strengthen their argument. They concluded that Deltadromeus should be regarded as a junior synonym of Bahariasaurus. The results of their phylogenetic analysis are displayed in the cladogram below, with Bahariasaurus (including Deltadromeus) indicated in the so-called "abelisauroid clade 1". |label1=Abelisauroidea}} In a paper analyzing the relationships of Mirischia and Santanaraptor, two coelurosaurs from the Cretaceous of Brazil, Brazilian paleontologist Rafael Delcourt and colleagues found Deltadromeus and Gualicho to be early-branching ornithomimosaurs in their phylogenetic analysis. Delcourt and colleagues (2025) noted that Deltadromeus' femur lacks a crest seen in many other theropods, a condition observed in ornithomimids like Struthiomimus and Gallimimus. Additionally, the medial condyle of the tibia of Deltadromeus bears a large process found in the tibiae of ornithomimosaurs like Mirischia. Although several African theropods from Gondwana have been theorized to be ornithomimosaurs, many of these taxa have since been reclassified as noasaurids, which too have edentulousness, gracile limbs, and elongated (neck) vertebrae like the ornithomimosaurs. However, Delcourt and colleagues as well as a conference abstract recovered Deltadromeus and Gualicho within Ornithomimosauria in their phylogenetic analyses, with the former paper stating that the classification of the two needs further study. == Paleoenvironment ==
Paleoenvironment
, with a group of Bahariasaurus in the far right background Fossils of Deltadromeus are known from the Kem Kem Beds, one of many dinosaur-bearing Cretaceous-aged geologic units in North Africa. Others include the Bahariya Formation of Egypt and the Elrhaz and Echkar Formations of Niger. The composition of the dinosaur fauna of North Africa at this time is an anomaly, as there are fewer herbivorous dinosaur species relative to carnivorous dinosaur species than in most fossil sites. This abundance of theropods compared to that of non-theropods was dubbed "Stromer's Riddle", which, despite suggestions that this is due to ecological, preservation, or other biases, can be supported by the fossil record. Isotopic evidence supports this as there were greater quantities of sizable, terrestrial animals in the diets of carcharodontosaurids and ceratosaurs from both the Kem Kem Beds and Elrhaz Formation. North Africa was dominated by a triumvirate of Abelisauroidea, Spinosauridae, and Carcharodontosauridae during the mid-Cretaceous, with all of these groups present in the Kem Kem Beds, Echkar, Elrhaz, and Bahariya Formations. which were recovered as a relative of Deltadromeus and Bahariasaurus in phylogenetic analyses. This would indicate niche partitioning in the large theropods in these localities, with Deltadromeus and Bahariasaurus filling a niche typically filled by ornithischians. However, this cannot be confirmed no skull material has been found for either Deltadromeus or Bahariasaurus. This river system was freshwater based on the presence of lungfishes and other typically freshwater vertebrates. This indicates that the Kem Kem Beds had a wide variety of features, including river channels, river banks and sandbars. an indeterminate somphospondylian titanosauriform, and an indeterminate titanosaur, one comparable in size to the giant Paralititan. Ornithischian fossils are extremely rare, only being represented from an isolated thyreophoran tooth As for theropods, many are known, including one or two distinct indeterminate abelisaurids, the carcharodontosaurids Carcharodontosaurus However, many of these dinosaurs are known from isolated or incomplete remains, have complicated taxonomies, or are under study. ==References==
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