Beaks and radulae of Early Carboniferous goniatites

About one hundred goniatite beaks (jaws) and five radulae from the Late Mississippian (Early Carboniferous) of Arkansas wgere studied with light and scanning electron microscopy (SEM). Four beaks were found within the body chamber of the goniatite Girtyoceras. Owing to the three-dimensional preservation, these oldest known beaks could be studied in detail and compared with those in living coleoids. The beaks are univalved, and the lower one is larger than the upper. Each beak consists of an organic outer and inner lamella; only the rostrum is weakly calcified. In the lower beak the outer and inner lamellae are about the same length, but in the upper beak the outer lamella is considerably shorter than the inner lamella. The goniatite beaks resemble those in living coleoids in the relative length of the outer and inner lamellae in the upper beak, which probably indicates similarity in muscle insertion. Concerning the length of the inner and outer lamellae, the lower beak is similar to that in Vampyroteuthis and the pelagic octopod Tremoctopus. Late Mississippian goniatite beaks dealt with here are similar to those of Carboniferous and Permian goniatites in general morphology, but differ from those of Mesozoic ammonoids. In the latter ammonoids, the lower beak has a long outer lamella and a short inner lamella, whereas both lamellae have about equal length in the goniatites. Goniatite radulae remain stable during ammonoid evolution and demonstrate a more or less distinct similarity with those in living coleoids.