Venaticosuchus

Fossils of Venaticosuchus have only been recovered, in the early sixties, from a single fossil site in Argentina. This site, located at Cerro Las Lajas in the province of La Rioja, contains outcrops from the middle part of the Ischigualasto Formation, which was deposited in the Late Triassic. The Cerro Las Lajas site has also preserved fossils of Trialestes (one of the oldest crocodylomorphs) and Pisanosaurus (either a silesaurid or one of the oldest ornithischian dinosaurs). However, its overall amount of fossil material is scant relative to other Ischigualasto localities, and few fossils found at Cerro Las Lajas have been found elsewhere in the formation.[1] This makes it difficult to gauge exactly when the Cerro Las Lajas site was formed, but it had to have been between ~231.7 and ~225.0 million years ago, based on radiometric dating of the youngest and oldest rocks in the Ischigualasto Formation.[3]

Venaticosuchus fossil remains consist of skull and jaw material from a single animal, although much of the braincase and the top part of the skull are missing. The original 1970/1971 description of Venaticosuchus, published by renowned Argentine paleontologist José Bonaparte, derived the generic name from the Latin venaticus, "hunting", and the Latinised Greek suchus, "crocodile". The specific name honoured the paleontologist Carlos Rusconi Sassi, deceased in 1969. It indicated that the fossil also included a partial forelimb and osteoderms (bony scutes),[4] but these remains have not been located by modern paleontologists and are thus considered missing.[5] Bones from the right side of the skull were also considered missing until 2015, when quadratojugal, quadrate, surangular, articular, and angular bones from the right side of the skull were rediscovered. These were described in 2018 as part of a study on ornithosuchid jaw biomechanics.[2] All known fossil material of Venaticosuchus is stored in the vertebrate paleontology collections of the Instituto Miguel Lillo at the National University of Tucumán. There, they are collectively termed the holotype specimen PVL 2578.[1]

Life restoration

The total body length of Venaticosuchus is unknown, considering that only skull and jaw material is known in the present day. The jaw had a total length of 26.0 centimeters (10.2 inches), which was marginally larger than the largest known jaws from Ornithosuchus and Riojasuchus (excluding the massive and enigmatic archosaur "Dasygnathoides", which has historically been considered a synonym of Ornithosuchus).[6] Other ornithosuchids were medium-sized reptiles, slightly less than 2 meters (6.6 feet) in length.[2]

The side of the snout was formed by the toothed maxilla and premaxilla bones, which were separated by a toothless gap known as a diastema. This is similar to the case in other ornithosuchids, and Venaticosuchus particularly resembles Riojasuchus due to the maxilla curving upwards behind the diastema and the premaxilla hooking downwards in front of it. The premaxilla is not preserved well enough to conclude anything about its tooth count, but other ornithosuchids had three premaxillary teeth. The maxilla in general resembles that of Riojasuchus, with a triangular antorbital fenestra with a tapering front tip. On the other hand, the antorbital fossa (the basin in which the antorbital fenestra lies) is shallower, smaller, and more smoothly textured than that of Riojasuchus. There were eight teeth in the maxilla; these teeth were large and serrated, with the second tooth being the largest. They were teardrop-shaped in cross section, unlike other ornithosuchids. Similar to Riojasuchus and the erpetosuchid Pagosvenator[7], the jugal (cheek bone) had a Y-shaped ascending process (upward-branching bone) which defined the lower edge of the eye socket.[1] The quadratojugal bone (at the rear lower corner of the skull) was L-shaped, with two branches converging at a 45 degree angle. It was also smoother than that of other ornithosuchids, in which there were ridges on the two branches of the quadratojugal.[2] The quadrate bone, which lies adjacent to the quadratojugal and forms the upper half of the jaw joint), forms a steep angle identical to that of the quadratojugal. The quadrate is also smooth, and it encloses a circular hole (only visible from behind) along its contact with the quadratojugal.[2]

Part of the nasal cavity was filled in by sediment, showing that the olfactory bulbs were separated at the midline. The braincase is incomplete, only preserving the parabasisphenoid (the bone which forms the base of the braincase). As with other archosaurs, the rear lower part of the braincase possessed two different pairs of large, rounded plates: The basipterygoid processes (which are positioned at the base of the braincase and contact the roof of the mouth) and the basitubera (which are positioned further up and do not contact any bone, acting as levers for the musculi recti capitis anteriores that make the head nod). The basipterygoid processes are separated from the basitubera by means of a pronounced notch which forms an acute angle (~60 degrees). This angle is more acute than any other early pseudosuchian (for example, Riojasuchus's is about 82 degrees), so it can be considered a unique feature of Venaticosuchus. The basipterygoid processes themselves were also thinner than those of Riojasuchus, and they are separated from each other by a wider gap. The palate (roof of the mouth) was generally similar to that of Riojasuchus.[1]

The mandible (lower jaw) was robust, proportionally more similar to Ornithosuchus rather than Riojasuchus. On the other hand, the mandibular fenestra (a large hole on the side of the jaw) was elongated and proportionally more similar to that of Riojasuchus rather than Ornithosuchus. The dentary (the main toothed bone of the mandible) had a rounded front end which is also expanded upwards. The first two teeth of the dentary were very large, overlapping the snout at the level of the maxillary/premaxillary diastema. This contrasts with other ornithosuchids, which have a smaller tooth in front of the two enlarged teeth.[2] The two enlarged dentary teeth of Venaticosuchus were unserrated and oval-shaped in cross section. Most of the other mandibular teeth could not be assessed due to the overlapping snout obscuring details, but seem to resemble the maxillary teeth. The surangular and articular bones (which form the rear upper part of the mandible) were smooth and thin. These features are unique to Venaticosuchus compared to other ornithosuchids, which have a noticeable pit on both the outer surface of the surangular and the inner surface of the articular.[1][2] The angular bone (which forms the rear lower part of the mandible) was elongated, forming the entire lower edge of the mandibular fenestra and being covered with striations. Both of these features are also in contrast with other ornithosuchids.[1]