Parareptilia


Parareptilia ("at the side of reptiles") is a subclass or clade of basal sauropsids (reptiles), typically considered the sister taxon to Eureptilia (the group that likely contains all living reptiles). Parareptiles first arose near the end of the Carboniferous period and achieved their highest diversity during the Permian period. Several ecological innovations were first accomplished by parareptiles among reptiles. These include the first reptiles to return to marine ecosystems (mesosaurs), the first bipedal reptiles (bolosaurids such as Eudibamus), the first reptiles with advanced hearing systems (nycteroleterids and others), and the first large herbivorous reptiles (the pareiasaurs). The only parareptiles to survive into the Triassic period were the procolophonoids, a group of small generalists, omnivores, and herbivores. The largest family of procolophonoids, the procolophonids, rediversified in the Triassic and continued to be successful until their demise in the Triassic-Jurassic mass extinction.[2][3]

Compared to most eureptiles, parareptiles retained fairly "primitive" characteristics such as robust, low-slung bodies and large supratemporal bones at the back of the skull. While all but the earliest eureptiles were diapsids, with two openings at the back of the skull, parareptiles were generally more conservative in the extent of temporal fenestration. In its modern usage, Parareptilia was first utilized as a cladistically correct alternative to Anapsida, a term which historically referred to reptiles with solid skulls lacking holes behind the eyes.[4] Nevertheless, not all parareptiles have 'anapsid' skulls, and some do have large holes in the back of the skull. They also had several unique adaptations, such as a large pit on the maxilla, a broad prefrontal-palatine contact, and the absence of a supraglenoid foramen of the scapula.[4][5]

Like many other so-called 'anapsids', parareptiles were historically understudied. Interest in their relationships were reinvigorated in the 1990s, when several studies argued that Testudines (turtles and their kin) were members of Parareptilia.[4] Although this would suggest that Parareptilia was not extinct after all, the origin of turtles is still heavily debated. Many other morphological or genetic analyses find more support for turtles among diapsid eureptiles such as sauropterygians or archosauromorphs, rather than parareptiles.[6][7][8][3]

Parareptilian skulls were diverse, from mesosaurs with elongated snouts filled with hundreds of thin teeth, to the snub-nosed, knob-encrusted skulls of pareiasaurs. Parareptile teeth were quite variable in shape and function between different species. However, they were relatively homogenous on the same skull. While most synapsids and many early eureptiles had a caniform region of enlarged fang-like teeth in the front half of the skull, very few parareptiles possessed caniform teeth.[5]

Many amniotes have a row of small pits running along bones at the edge of the mouth, but parareptiles have only a few pits, with one especially large pit near the front of the maxilla.[4][9][7] The rest of the skull was often strongly-textured by pits, ridges, and rugosities in most parareptile groups, occasionally culminating in complex bosses or spines. The maxilla is usually low, while the prefrontal and lacrimal bones in front of the eye are both fairly large. In all parareptiles except mesosaurs, the prefrontal has a plate-like inner branch which forms a broad contact with the palatine bone of the palate.[4][7][5] A prominent hole, the foramen orbitonasale, is present at the intersection of the prefrontal, palatine, and lacrimal. Parareptilian palates also have toothless and reduced ectopterygoid bones, a condition taken to extremes in mesosaurs, which have lost the ectopterygoid entirely.[4][5]


A skull diagram of Colobomycter pholeter, a probable acleistorhinid from the Early Permian. Unlike most parareptiles, this species lacks tabular bones and has a heterodont dentition with large caniform teeth.
A skeletal diagram of Sauropareion anoplus, a procolophonid from the Early Triassic