Article: The braincase and jaws of Cladodus from the Lower Carboniferous of Scotland

The jaws and an allegedly associated orbitotemporal region of a shark braincase are described from the Lower Carboniferous (Upper Visean, Brigantian Stage, Clackmannan Group, Lower Limestone Formation) of Lugton, Ayrshire, Scotland. The braincase specimen is important because it is associated with a tooth of Cladodus elegans Newberry and Worthen, 1870, a form considered to be close to the type species Cladodus mirabilis Agassiz, 1843. An occipital fragment, a basicranium and a fragment of palatoquadrate from the same locality are also described. All the material probably represents the same species, and it is even possible that some of the material came from a single individual. The braincase resembles those of Cladodoides and Tamiobatis, especially in the proportions of its otic region, the presence of an extensive dorsum sellae, and the arrangement of canals for major nerves and blood vessels. The internal structure of the occipital region is remarkably like that found in Tamiobatis braincases in having a prominent subnotochordal septum and paired subnotochordal and paroccipital chambers. Cladodus is no longer regarded as a nomen dubium, since only a few species founded on isolated teeth are now retained in the re-diagnosed genus, and one of these (C. elegans) is now associated with skeletal remains. The best systematic placement for Cladodus sensu stricto is among the Ctenacanthiformes or even the Ctenacanthidae. Cladodus may be a synonym of Ctenacanthus, but this cannot be confirmed with the available material.