Deltadromeus

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Deltadromeus
Temporal range: Late Cretaceous (Cenomanian), 95  Ma
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Deltadromeus in Japan.jpg
Mounted skeleton cast with reconstructed skull
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Noasauridae (?)
Genus: Deltadromeus
Sereno et al., 1996
Species:
D. agilis
Binomial name
Deltadromeus agilis
Sereno et al., 1996
Synonyms

Deltadromeus (meaning "delta runner") is a genus of theropod dinosaur from the Aoufous Formation of Morocco. [1]

It had long, unusually slender hind limbs for its size, suggesting that it was a swift runner. [1] The skull is not known. One fossil specimen of a single species (D. agilis, or "agile delta runner") has been described, found in the Kem Kem Beds, which date to the mid-Cretaceous Period (mid-Cenomanian age), about 95 million years ago. It may be a junior synonym of the contemporary Bahariasaurus . [2] The classification of Deltadromeus has been in flux since its original description. In 2016, a South American theropod known as Gualicho shinyae was found to possess many similarities with Deltadromeus. Depending on the phylogenetic position of Gualicho, Deltadromeus may have been a neovenatorid carnosaur, a tyrannosauroid, or a basal coelurosaur if its close relation to Gualicho is legitimate. [3] [4] [5] Other studies have considered it a ceratosaurian, more specifically a member of the family Noasauridae. [6] [7] [8] [9]

Description

Deltadromeus feeding on a sauropod Deltadromeus31DB.jpg
Deltadromeus feeding on a sauropod

The holotype of Deltadromeus agilis (museum catalogue number SGM-Din2) is a partial skeleton of an animal which is estimated to have measured 8 metres (26 ft) long. The weight of the living animal was estimated to have been around 1,050 kilograms, slightly more than an imperial ton. [10]

Size of the holotype, speculatively restored as a noasaurid, compared to a human Deltadromeus Scale.svg
Size of the holotype, speculatively restored as a noasaurid, compared to a human

A number of specimens (catalogued under IPHG 1912 VIII) were originally considered by Ernst Stromer to be conspecific with Bahariasaurus , [11] but were referred to Deltadromeus by Paul Sereno in 1996. [1] They were thought to come from a much larger individual, with a femur (upper leg bone) length of 1.22 metres (4 ft), compared to the 0.74-metre-long (2.4 ft) femur of the holotype. These referred specimens, if legitimately assigned to Deltadromeus, would have indicated that members of the genus could grow up to 12.2 metres (40 ft) in length, approximately the size of a Tyrannosaurus rex . [1] However, the referral of the coracoid, pubes and hindlimb material catalogued under IPHG 1912 VIII to Deltadromeus has been questioned because the remains came from different horizons and localities in the Bahariya Formation, and actually exhibit notable differences from the holotype of Deltadromeus. [12]

The Deltadromeus skeleton has been found in the same formation as those of the giant theropods Carcharodontosaurus , Spinosaurus and Bahariasaurus, which may be synonymous with Deltadromeus. No skull material has been found for either Deltadromeus or Bahariasaurus, and though carnivore teeth labelled as "Deltadromeus" are sold in rock shops, there is no way of knowing if they actually come from this animal. [2]

Classification

Diagrams showing holotype remains (A) and size of a lost femur (B) Deltadromeus skeletal.jpg
Diagrams showing holotype remains (A) and size of a lost femur (B)
Pectoral and forelimb Deltadromeus forelimb.jpg
Pectoral and forelimb

As a ceratosaur

Many studies published since the original description of Deltadromeus have considered it to be a ceratosaur, although different studies disagree on what kind of ceratosaur. One 2003 study suggested it was a member of the Noasauridae, [6] though others have found it to be more primitive, possibly related to the primitive ceratosaurs Elaphrosaurus and Limusaurus . [13] [14] A more comprehensive study of noasaurid relationships published in 2016 found that both of these interpretations were essentially correct, with Deltadromeus, Limusaurus and Elaphrosaurus all found to be within the Noasauridae. [7] 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 writers of the paper, resolving the phylogenetic positions of Gualicho, Aoniraptor, Deltadromeus and megaraptorans is one of the most critical issues presently facing theropod systematics. [8] Deltadromeus was also considered a noasaurid in a 2020 review of the Kem Kem Group geology and fauna. [9]

The cladogram below follows a 2016 analysis by Oliver Rauhut and Matthew Carrano. [7]

Abelisauroidea  

As an avetheropod

The original description of Deltadromeus in 1996 found that it was a fairly basal coelurosaur, only slightly more advanced than the Late Jurassic genus Ornitholestes . [1]

Cast with alternate skull reconstruction Deltadromeus agilis (2).jpg
Cast with alternate skull reconstruction

In 2016, an analysis of Gualicho , a South American theropod considered to belong to the allosauroid family Neovenatoridae, found Deltadromeus to be Gualicho's probable sister taxon. However, the analysis also noted that Deltadromeus shared many features with ceratosaurs and that if Gualicho was removed from the analysis, Deltadromeus would resolve to a member of Ceratosauria. [3] In an analysis of Aoniraptor , which may be the same animal as Gualicho, Deltadromeus was found along with Aoniraptor and Bahariasaurus to probably form a still poorly known clade of megaraptoran tyrannosauroids different from the Megaraptoridae. [4] A 2018 study by Porfiri et al. has supported the idea that Gualicho and megaraptorans were basal coelurosaurs, outside of both Neovenatoridae and Tyrannosauroidea. However, this study did not include Deltadromeus. [5]

The cladogram below follows the 2016 Gualicho analysis by Sebastián Apesteguía, Nathan D. Smith, Rubén Juarez Valieri and Peter J. Makovicky. [3]

Related Research Articles

<i>Carcharodontosaurus</i> Genus of carcharodontosaurid dinosaur from the Cretaceous period

Carcharodontosaurus is a genus of carnivorous theropod dinosaur that lived in North Africa from about 99 to 94 million years ago during the Cenomanian stage of the Late Cretaceous. Two teeth of the genus, now lost, were first described from Algeria by French paleontologists Charles Depéret and Justin Savornin as Megalosaurus saharicus. A partial skeleton was collected by crews of German paleontologist Ernst Stromer during a 1914 expedition to Egypt. Stromer did not report the Egyptian find until 1931, in which he dubbed the novel genus Carcharodontosaurus, making the type species C. saharicus. Unfortunately, this skeleton was destroyed during the Second World War. In 1995 a nearly complete skull of C. saharicus, the first well-preserved specimen to be found in almost a century, was discovered in the Kem Kem Beds of Morocco; it was designated the neotype in 1996. Fossils unearthed from the Echkar Formation of northern Niger were described and named as another species, C. iguidensis, in 2007.

<span class="mw-page-title-main">Ceratosauria</span> Extinct clade of dinosaurs

Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with Ceratosaurus than with birds. The oldest known ceratosaur, Saltriovenator, dates to the earliest part of the Jurassic, around 199 million years ago. Ceratosauria includes three major clades: Ceratosauridae, Noasauridae, and Abelisauridae, found primarily in the Southern Hemisphere. Originally, Ceratosauria included the above dinosaurs plus the Late Triassic to Early Jurassic Coelophysoidea and Dilophosauridae, implying a much earlier divergence of ceratosaurs from other theropods. However, most recent studies have shown that coelophysoids and dilophosaurids do not form a natural group with other ceratosaurs, and are excluded from this group.

<i>Masiakasaurus</i> Noasaurid theropod dinosaur genus from the Late Cretaceous period

Masiakasaurus is a genus of small predatory noasaurid theropod dinosaurs from the Late Cretaceous of Madagascar. In Malagasy, masiaka means "vicious"; thus, the genus name means "vicious lizard". The type species, Masiakasaurus knopfleri, was named after the musician Mark Knopfler, whose music inspired the expedition crew. It was named in 2001 by Scott D. Sampson, Matthew Carrano, and Catherine A. Forster. Unlike most theropods, the front teeth of M. knopfleri projected forward instead of straight down. This unique dentition suggests that they had a specialized diet, perhaps including fish and other small prey. Other bones of the skeleton indicate that Masiakasaurus were bipedal, with much shorter forelimbs than hindlimbs. M. knopfleri was a small theropod, reaching 1.8–2.1 m (5.9–6.9 ft) long and weighing 20 kg (44 lb).

<i>Bahariasaurus</i> Extinct genus of dinosaurs

Bahariasaurus is an enigmatic genus of large theropod dinosaur. Bahariasaurus is known to have included at least 1 species, Bahariasaurus ingens, which was found in North African rock layers dating to the Cenomanian and Turonian ages of the Late Cretaceous. The only fossils confidently assigned to Bahariasaurus were found in the Bahariya Formation of the Bahariya oasis in Egypt by Ernst Stromer but were destroyed during a World War II bombing raid with the same raid taking out the holotype of Spinosaurus and Aegyptosaurus among other animals found in the Bahariya Formation. While there have been more fossils assigned to the genus such as some from the Farak Formation of Niger, these remains are referred to with much less certainty. Bahariasaurus is, by most estimations, one of the largest theropods, approaching the height and length of other large bodied theropods such as Tyrannosaurus rex and the contemporaneous Carcharodontosaurus. The aforementioned estimations tend to put it at around 11–12.2 metres in length and 4-4.8 tonnes in overall weight.

<i>Chilantaisaurus</i> Theropod dinosaur genus from the Late Cretaceous period

Chilantaisaurus is a genus of large theropod dinosaur, possibly a neovenatorid or a primitive coelurosaur, from the Late Cretaceous Ulansuhai Formation of China. The type species, C. tashuikouensis, was described by Hu in 1964.

<span class="mw-page-title-main">Abelisauroidea</span> Extinct clade of dinosaurs

Abelisauroidea is a diverse superfamily of ceratosaurian dinosaurs, typically regarded as a Cretaceous group, though the earliest abelisaurid remains are known from the Middle Jurassic of Argentina and possibly Madagascar. Possible Abelisauridae remains were also discovered in Late Jurassic Tendaguru Beds in Tanzania.

<i>Elaphrosaurus</i> Ceratosaurian theropod dinosaur genus from the Late Jurassic Period

Elaphrosaurus is a genus of ceratosaurian theropod dinosaur that lived approximately 154 to 150 million years ago during the Late Jurassic Period in what is now Tanzania in Africa. Elaphrosaurus was a medium-sized but lightly built member of the group that could grow up to 6.2 m (20 ft) long. Morphologically, this dinosaur is significant in two ways. Firstly, it has a relatively long body but is very shallow-chested for a theropod of its size. Secondly, it has very short hindlimbs in comparison with its body. Phylogenetic analyses indicate that this genus is likely a ceratosaur. Earlier suggestions that it is a late surviving coelophysoid have been examined but generally dismissed. Elaphrosaurus is currently believed to be a very close relative of Limusaurus, an unusual beaked ceratosaurian which may have been either herbivorous or omnivorous.

<i>Megaraptor</i> Extinct genus of dinosaurs

Megaraptor is a genus of large theropod dinosaur that lived in the ages of the Late Cretaceous. Its fossils have been discovered in the Patagonian Portezuelo Formation of Argentina, South America. Initially thought to have been a giant dromaeosaur-like coelurosaur, it was classified as a neovenatorid allosauroid in previous phylogenies, but more recent phylogeny and discoveries of related megaraptoran genera has placed it as either a basal tyrannosauroid or a basal coelurosaur with some studies still considering it a neovenatorid.

<i>Noasaurus</i> Extinct genus of dinosaurs

Noasaurus is a genus of ceratosaurian theropod dinosaur genus from the late Campanian-Maastrichtian of Argentina. The type and only species is N. leali.

<i>Spinostropheus</i> Extinct genus of dinosaurs

Spinostropheus is a genus of carnivorous neotheropod theropod dinosaur that lived in the Middle Jurassic period and has been found in the Tiouraren Formation, Niger. The type and only species is S. gautieri.

<span class="mw-page-title-main">Noasauridae</span> Extinct family of dinosaurs

Noasauridae is an extinct family of theropod dinosaurs belonging to the group Ceratosauria. They were closely related to the short-armed abelisaurids, although most noasaurids had much more traditional body types generally similar to other theropods. Their heads, on the other hand, had unusual adaptations depending on the subfamily. 'Traditional' noasaurids, sometimes grouped in the subfamily Noasaurinae, had sharp teeth which splayed outwards from a downturned lower jaw.

<i>Orkoraptor</i> Extinct genus of dinosaurs

Orkoraptor is a genus of medium-sized megaraptoran theropod dinosaur from the late Cretaceous Period of Argentina. It is known from incomplete fossil remains including parts of the skull, teeth, tail vertebrae, and a partial tibia. The specialized teeth resemble those of some maniraptoriform theropods, namely the deinonychosaurs and compsognathids. This and other anatomical features led the authors who described it to suggest that it was a maniraptoran coelurosaur. However, subsequent studies found it to be a megaraptoran. Found in the Cerro Fortaleza Formation of Southern Patagonia, it is one of the southernmost carnivorous dinosaurs known from South America.

<i>Aerosteon</i> Extinct genus of dinosaurs

Aerosteon is a genus of megaraptoran dinosaur from the Late Cretaceous period of Argentina. Its remains were discovered in 1996 in the Anacleto Formation, which is from the late Campanian. The type and only known species is A. riocoloradensis. Its specific name indicates that its remains were found 1 km north of the Río Colorado, in Mendoza Province, Argentina.

<span class="mw-page-title-main">Bahariasauridae</span> Probable family of averostran theropods

Bahariasauridae is a potential family of averostran theropods that might include a handful of African and South American genera, such as Aoniraptor, Bahariasaurus, Deltadromeus, and Gualicho. The placement of these theropods is controversial, with some studies placing them as basal ceratosaurs possibly related to Noasauridae, others classifying them as megaraptorans, basal neovenatorids, or basal coelurosaurs. There is also a possibility the group might not be monophyletic, as a monograph on the vertebrate diversity in the Kem Kem Beds published in 2020 found Bahariasaurus to be nomen dubium. In the same paper Deltadromeus is classified as an noasaurid, a result also recovered by some previous studies. A 2024 analysis found Aoniraptor, Bahariasaurus, Deltadromeus, Elaphrosaurus and Gualicho to form a monophyletic clade at the base of Ceratosauria.

<i>Limusaurus</i> Genus of theropod dinosaur

Limusaurus is a genus of theropod dinosaur that lived in what is now China during the Late Jurassic, around 161 to 157 million years ago. The type and only species Limusaurus inextricabilis was described in 2009 from specimens found in the Upper Shishugou Formation in the Junggar Basin of China. The genus name consists of the Latin words for "mud" and "lizard", and the species name means "impossible to extricate", both referring to these specimens possibly dying after being mired. Limusaurus was a small, slender animal, about 1.7 m in length and 15 kg (33 lb) in weight, which had a long neck and legs but very small forelimbs. It underwent a drastic morphological transformation as it aged: while juveniles were toothed, these teeth were completely lost and replaced by a beak with age. Several of these features were convergently similar to the later ornithomimid theropods as well as the earlier non-dinosaurian shuvosaurids.

<span class="mw-page-title-main">Neovenatoridae</span> Extinct family of dinosaurs

Neovenatoridae is a proposed clade of carcharodontosaurian dinosaurs uniting some primitive members of the group such as Neovenator with the Megaraptora, a group of theropods with controversial affinities. Other studies recover megaraptorans as basal coelurosaurs unrelated to carcharodontosaurs. Other theropods with uncertain affinities such as Gualicho, Chilantaisaurus and Deltadromeus are also sometimes included.

<span class="mw-page-title-main">Megaraptora</span> Extinct clade of dinosaurs

Megaraptora is a clade of carnivorous theropod dinosaurs. Its derived members, the Megaraptoridae are noted for their large hand claws and powerfully-built forelimbs, which are usually reduced in size in other large theropods.

<span class="mw-page-title-main">Timeline of ceratosaur research</span>

This timeline of ceratosaur research is a chronological listing of events in the history of paleontology focused on the ceratosaurs, a group of relatively primitive, often horned, predatory theropod dinosaurs that became the apex predators of the southern hemisphere during the Late Cretaceous. The nature and taxonomic composition of the Ceratosauria has been controversial since the group was first distinguished in the late 19th century. In 1884 Othniel Charles Marsh described the new genus and species Ceratosaurus nasicornis from the Late Jurassic Morrison Formation of the western United States. He felt that it belonged in a new family that he called the Ceratosauridae. He created the new taxon Ceratosauria to include both the Ceratosauridae and the ostrich-like ornithomimids. The idea of the Ceratosauria was soon contested, however. Later that same decade both Lydekker and Marsh's hated rival Edward Drinker Cope argued that the taxon was invalid.

Gualicho is a genus of theropod dinosaur. The type species is Gualicho shinyae. It lived in what is now northern Patagonia, on what was then a South American island continent split off from the supercontinent Gondwana. The fossils were found in the Huincul Formation, dating to the late Cenomanian-early Turonian age of the upper Cretaceous Period, around 91 million years ago.

<i>Afromimus</i> Extinct genus of dinosaurs

Afromimus is a genus of 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, vertebrae, and ribs found in the Ténéré Desert. It was originally classified as an ornithomimosaurian, but subsequently it was argued to be an abelisauroid.

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