Terrestrisuchus

Size comparison

Terrestrisuchus was a small, slender crocodylomorph with very long legs, quite unlike modern crocodilians. It was initially estimated to have been between 49 to 76 centimetres (19 to 30 in) long, although this estimate may be based on juvenile specimens and fully grown Terrestrisuchus may have reached or exceeded 1 metre (3.3 ft) in length.[1][2]

Its skull was long and narrow, with a tapering, pointed triangular snout lined with sharp curved teeth. The upper jaw margin was straight, and lacked a diastema (a gap in the tooth row) between the maxilla and the premaxilla. By contrast, the long and slender dentary bones of the lower jaw curved slightly upwards towards the front.[1] Unlike modern crocodilians, the eye of Terrestrisuchus was supported by a ring of bony ossicles, the sclerotic ring.[3]

Skeletal restoration showing known remains

The body was relatively short and shallow, and the spine was topped by paired rows of osteoderms running down from its neck down its back. These osteoderms are described as "leaf-shaped", being relatively longer than wide with a prominent spur at the front that slides under and interlocks with the scute in front of it. This provides a rigid support for the body and limited the flexibility of its spine, supporting its body on land.[4] The hips of Terrestrisuchus had an elongated pubis, unlike living crocodilians. Terrestrisuchus is also known to have had tightly packed gastralia, or belly ribs. Its tail was particularly long, about twice the length of the head and body combined with an estimated 70 caudal vertebrae in total, and may have been used as a balance allowing the animal to rear up and run on its hind legs for brief periods.[1][5]

The first fossils of Terrestrisuchus were discovered by Professor K. A. Kermack and Dr. P. L. Robinson in the spring of 1952, recovered from the Pant-y-ffynon Quarry located near Cowbridhe, Glamorgan in South Wales. Their finds were presented by Kermack to the Linnean Society of London on October 8, 1953, and was recognised belonging to a "primitive crocodile or crocodile ancestor". No osteoderms had been identified yet at the time, which Kermack regarded as representing a "missing link" between modern crocodilians and the Triassic "thecodonts". The fossils included several well-preserved articulated partial skeletons and various isolated bones. Kermack refrained from naming the animal or nominating a type specimen, as preparation of the fossils was still ongoing. The specimens were eventually named and thoroughly described by P. J. Crush in 1984, with the generic name Terrestrisuchus chosen to emphasise the terrestrial lifestyle of this crocodylomorph, and the specific name from the Latin gracilis for its light, graceful build.[1][12]

The Pant-y-ffynnon Quarry is composed mostly of Carboniferous limestone, but the fossils of Terrestrisuchus were recovered from Triassic sedimentary rocks that were deposited within fissures in the limestone (such as sandstones and marls). The age of the deposits has been historically debated, with older literature suggesting a Carnian to Norian age. However, palynological data has been used to determine a younger Rhaetian age, close to the very end of the Triassic. This estimate has been corroborated by Rhaetian index fossils such as conchostracans and geomorphological data.[13]

Additional material attributed to Terrestrisuchus has been discovered in other Late Triassic fissure deposits in South Wales and Bristol, including the Ruthin Quarry in Wales and the Tytherington and Cromhall quarries near Bristol, as well as a possible specimen from Durdham Down.[14][15] The fossils of Terrestrisuchus were originally housed at University College, London before being transferred to the Natural History Museum in London where they are currently stored.[16]

Terrestrisuchus was originally classified as a member of the crocodylomorph suborder "Sphenosuchia", a group that included various other similar long-legged early crocodylomorphs and was considered to be a separate radiation from the group that all later crocodylomorphs would evolve from.[1] However, this classification was made prior to the invention of cladistic phylogenetic analyses, which has since demonstrated that "Sphenosuchia" is an unnatural grouping (paraphyletic), meaning that "sphenosuchians" are not all descended from a single common ancestor to the exclusion of all other crocodylomorphs. Instead, the "sphenosuchians" are a grade of basal crocodylomorphs that lead up to the more derived crocodyliforms.[16] Nonetheless, Terrestrisuchus has consistently been recovered within this grade, as shown in the cladogram below from Leardi et al. (2017):[17]

Terrestrisuchus depicted hunting a cockroach

The thin, serrated teeth of Terrestrisuchus indicate that it was carnivorous, and like other early crocodylomorphs it was likely a generalist pursuit hunter that preyed upon small to mid-sized prey items.[2][13] The shape and construction of its hip bones, particularly the elongated pubis, indicate that they were rigidly sutured together and that its pubis was not mobile as it is in modern crocodilians. This indicates that Terrestrisuchus would not have utilised the hepatic piston method of breathing found in modern crocodilians.[22]

Terrestrisuchus was a coastal species, unusual for basal crocodylomorphs, which are typically known from inland floodplain environments. In the Late Triassic, the Pant-y-ffynnon Quarry and other quarries were part of ancient islands in a palaeo-archipelago that stretched across southern Wales and England to Bristol. The islands were forested karstic environments, riddled with fissures, sinkholes and caverns eroded into the limestone, environments which long limbed, agile reptiles like Terrestrisuchus may have been well suited to inhabit.[24]

On the Pant-y-ffynnon palaeo-island, Terrestrisuchus coexisted with other archosaurs such as the similarly long-legged, enigmatic pseudosuchian Aenigmaspina, the herbivorous sauropodomorph dinosaur Pantydraco, and possibly a coelophysoid theropod. Rhynchocephalians (relatives of modern tuataras) were very abundant and are known from at least three species including Clevosaurus cambrica, Diphydontosaurus and at least one or two unnamed species, which likely formed a large part of the diet of Terrestrisuchus (possibly evidenced by bite marks on the bones of Clevosaurus that likely belong to Terrestrisuchus). Terrestrisuchus was a relatively rare component of the island's fauna, an expected relationship for a predatory animal.[13]