Concavenator Skull Diagram

''Concavenator'' (meaning Cuenca hunter) is a genus of carcharodontosaurian dinosaur that lived in Spain during the Early Cretaceous epoch, about 125 million years ago. The genus contains a single species, ''Concavenator corcovatus'', named and described in 2010 from a nearly complete skeleton collected from the Las Hoyas fossil site of the La Huérguina Formation.

''Concavenator'' was a medium-sized carcharodontosaurian, reaching about in length and in weight. Unlike most carcharodontosaurians, the neural spines of the last dorsal (back) vertebrae were tall, creating a sail-like structure.

History

The first fossil remains of ''Concavenator'' were recovered from limestone slabs of Las Hoyas locality (considered a '' Konservat-Lagerstätten''; deposits of exceptional fossil preservation) of Cuenca Province, Spain, which belongs to La Huérguina Formation. The remains were represented by an articulated, nearly complete skeleton of a theropod dinosaur individual encased in limestone, comprising the skull, ten cervical vertebrae, thirteen dorsal vertebrae (with the last two presenting an unusual elongation), five sacral vertebrae, thirty caudal vertebrae, a partial pectoral girdle and arm, the pelvic girdle and partial legs, as well as ribs. This specimen was also found preserving integument traces, such as scale impressions on the feet and tail, which is characteristic of many of the lithographic limestones within La Huérguina Formation.

In 2010, the specimen was catalogued MCCM-LH 6666 and described by paleontologists Francisco Ortega, Fernando Escaso, and José Luis Sanz, used as the holotype for naming the new genus and species of carcharodontosaurid dinosaur ''Concavenator corcovatus''. The specimen belongs to the collection of the Science Museum of Castilla-La Mancha. The generic name, ''Concavenator'', is derived from the latin ''Conca'', in reference to the region of discovery, the Cuenca Province, and ''venator'', which means hunter. The specific name, ''corcovatus'', is taken from the latin ''corcovatus'', giving a glimpse onto the hump-like structure formed by the elongated dorsal vertebrae. In their description, the team noted that some anatomical elements had to be left unprepared (without removing the encasing rock), given the delicate nature of the preserved integument traces. Though initially described in 2010, a series of monographs about the taxon and holotype were published later, including examinations of the feet integument in 2015, as well as several anatomical redescriptions of the specimen in 2018.

Description

Size

''Concavenator'' was a medium-sized carcharodontosaurian, reaching long and . It possessed several unique features, including the two extremely tall vertebrae in front of the hips that formed a tall, narrow, pointed crest (possibly supporting a hump) on the dinosaur's back. The function of such crests, however, is currently unknown. Paleontologist Roger Benson from the University of Cambridge speculated that one possibility is that "it is analogous to head-crests used in visual displays", but the Spanish scientists who discovered it noted it could also be a thermal regulator.

Integument

''Concavenator'' had structures resembling quill knobs on its ulna, a feature known only in birds and other feathered theropods, such as dromaeosaurids. Quill knobs are created by ligaments which attach to the feather follicle and, since scales do not form from follicles, the authors ruled out the possibility that they could indicate the presence of long display scales on the arm. Instead, the knobs have been thought to probably anchor simple, hollow, quill-like structures. Such structures are known in both coelurosaurs, such as '' Dilong,'' and in some ornithischians, like ''Tianyulong'' and ''Psittacosaurus''. If the ornithischian quills are homologous with bird feathers, their presence in ''Concavenator'' and other allosauroids would be expected. However, if ornithischian quills are not related to feathers, the presence of these structures in ''Concavenator'' would show that feathers had begun to appear in earlier, more primitive forms than coelurosaurs.

Feathers or related structures would then likely be present in the first members of the clade Neotetanurae, which lived in the Middle Jurassic. No impressions of any kind of integument were found near the arm, although extensive scale impressions were preserved on other portions of the body, including broad, rectangular scales on the underside of the tail, bird-like scutes on the feet, and plantar pads on the undersides of the feet.

However, the significance of the 'quill knobs' remains controversial, as some amount of skepticism has been raised among experts on the validity of the interpretation that the ulnar bumps represent quill knobs. Christian Foth and colleagues noted that the quill knobs of ''Concavenator'' were on the anterolateral side of the ulna. They suggest they were intermuscular lines that acted as tendon attachments. The hypothesis that the bumps along the ulna represented muscular insertion points or ridges was subsequently examined and the results were presented at the 2015 meeting of the Society of Vertebrate Paleontology. Elena Cuesta Fidalgo, along with two of the researchers who initially described ''Concavenator'' (Ortega and Sanz), attempted to reconstruct its forearm musculature to determine if the ulnar bumps would be explained as an inter-muscular ridge. They identified the insertion point for the major arm muscles and determined that the row of bumps could not have been located between any of them. They found that the only possibility was that the bumps could be an attachment scar for the ''M. anconeus'' muscle. However, this is unlikely because this muscle normally attaches to a smooth surface without marks or bumps on the underlying bone. They argued that the most likely explanation for the bumps was their initial interpretation as feather quill knobs. The authors admitted that it was unusual for quill knobs to form along the posterolateral surface of the bone, but also noted that the same arrangement is found in some modern birds, like the Moorhen.Cuesta Fidalgo, Elena, Ortega, F., Sanz, J. (2015). Ulnar bumps of ''Concavenator'': Quill Knobs or Muscular scar? Myological Reconstruction of the forelimb of ''Concavenator corcovatus'' (Lower Cretaceous, Las Hoyas, Spain). ''Abstracts of papers of the 75th Annual Meeting of the Society of Vertebrate Paleontology'': 111-112.

In 2018, Cuesta Fidalgo published her doctorate thesis on the anatomy of ''Concavenator'', which argued that the ulna was preserved in lateral view. This means that the ulnar bumps were positioned posterolaterally instead of anterolaterally as Cau and Mortimer claimed. Cuesta Fidalgo noted that the proximal part of the ulna is affected by fracturing and abrasion, with certain features that would have shifted when compared to their position in the bone while the animal was alive. For example, in the fossil, the lateral process of the ulna is positioned further posteriorly than the ulnar bumps. In ''Allosaurus'' and ''Acrocanthosaurus'', the lateral process is on the lateral (rather than posterior) part of the bone, which would seem to support the ulnar bumps being anterolateral in position if the lateral process was truly preserved in lateral orientation in ''Concavenator''. However, Cuesta Fidalgo described how the lateral process was distorted posteriorly when compared to the bumps and was not valid evidence for the claim that the ulna had shifted into anterior view. The ulna's distortion (as well as genus-specific proportions) means that precise comparisons to ''Allosaurus'' and ''Acrocanthosaurus'' would be misleading. As Cuesta Fidalgo and her colleagues explained in 2015, the ulnar bumps could not be an intermuscular line if the bone is preserved in lateral view. Cuesta Fidalgo and her colleagues pointed out that these bumps on the ulna are posterolateral, which is unlike that of interosseous ligaments.

Classification

In their 2022 description of the giganotosaurin ''Meraxes'', ''Concavenator'' was recovered as an early-diverging member of the Carcharodontosauridae, in an unresolved polytomy with ''Eocarcharia'', ''Lajasvenator'', and ''Lusovenator''. The results of their phylogenetic analyses are displayed in the cladogram below:

See also

* ''Altispinax''
* ''Spinosaurus''
* List of animals with humps

References

External links

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{{Taxonbar, from=Q131715

Carcharodontosauridae
Dinosaur genera
Barremian dinosaurs
La Huérguina Formation
Fossil taxa described in 2010
Dinosaurs of Spain