Abstract
AbstractProcolophonids were diverse small reptiles through the late Permian and Triassic. Relatively complete specimens of various taxa are known from the Early and Late Triassic, but the 10 or so Middle Triassic taxa, from South Africa, Russia, China and the UK, are mostly incomplete, being known only from skulls or partial and poorly preserved isolated elements. Because of their small size, it has often been difficult to establish details of anatomy using physical preparation methods, so application of scanning technology can massively improve knowledge. Here, we describe the first substantial portion of a skull and anterior postcranial skeleton of the genus Kapes, known from Russia and the UK. CT scanning and 3D digital restoration of the new specimen from the Anisian Otter Sandstone of Devon, UK, reveal anatomical details previously unknown for this taxon, most notably the enlarged quadratojugal processes. This is also one of the few mature procolophonid specimens with a preserved postcranium, and it gives valuable insight into ossification patterns. Fused sutures of the skull, fused scapulocoracoid, and heavily worn teeth show evidence of the common reptilian ossification pattern with a possible limitation in tooth replacement. There is sufficient anatomical information to include Kapes bentoni for the first time in a cladistic analysis, which shows that it is sister to the Russian Kapes majmesculae and part of a clade of Anisian taxa phylogenetically more basal than Procolophon. Inferences on the function of the quadratojugal spines and fossorial mode of life are also considered.
Highlights
Procolophonids were diverse small reptiles through the late Permian and Triassic
That the partial postcranial skeleton of K. bentoni is known, we can address EXEMS 60/1985.9, a poorly preserved anterior portion of an interclavicle described by Milner et al (1990, fig. 7b) from the Otter Sandstone, and tentatively assigned by Spencer & Storrs (2002) to K. bentoni based on its size
The edge of the ridge is prominent in EXEMS 60/1985.9, whereas in BRSUG 29950-13 there is a gradual transition from the ridge to the rest of the surface of the lateral process
Summary
1968) from the Donguz Svita of the south Urals, of which there is an excellent skull housed in the Paleontological Institute, Moscow, PIN 4365/40, and described by Novikov & Sues (2004). Spencer & Storrs (2002) further discriminated Otter Sandstone procolophonids from other clades and presented some of their earlier jaw material as well as the holotype of a new species, Kapes bentoni, a partial snout, comprising the maxillae and partial premaxillae, palate and mandibles. This was an important discovery that has been widely accepted, but it was too incomplete to be coded for the seminal cladistic analysis of procolophonid phylogeny by Cisneros (2008a). The approximately 15 m thick Pennington Point Member is equivalent to Hounslow & McIntosh’s (2003) unit D which, on the basis of magnetostratigraphy, corresponds to the uppermost Anisian (Illyrian substage), possibly overlapping into the lowest Ladinian stage, making the age of these rocks c. 242 myr (Cohen et al 2013)
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