Late Cretaceous crocodile remains from Naskal (India): comparisons and biogeographic affinities

Crocodile teeth from the Maastrichtian inter-trappean beds of Naskal (peninsular India) are described here. Because of isolated denticles visible on sufficiently preserved carinae, the presence of a strong heterodonty (in size and shape), and by comparison to crocodile teeth from various taxa, they are considered as representing a ziphodont form with a heterodont dentition. The difference between ziphodont, "false ziphodont" and non-ziphodont dentitions is evaluated. With the help of scanning electron microscope photographs, it is shown that only precise characteristics of the denticles and not the tooth shape, allow to distinguish the three categories. These three categories do not correspond to monophyletic groups. It is also shown that the "alligatorid" heterodonty, meso- or eusuchian in grade, exists in each category. Although the ziphodont dentition is not sufficient to allow a taxonomical definition, the peculiarities that it often presents, depending on the taxa as well as the teeth shape, enable systematic approaches. An examination of previous works on the possible ziphodont crocodiles from the Tertiary deposits of the Indian subcontinent and on Naskal teeth demonstrate that the latter are closer to those of some Gondwanan crocodiles of mesosuchian grade, known from the early Cretaceous of Africa and possibly a form from the late Cretaceous of Madagascar. They are excluded from eusuchian Laurasiatic as well as Paleogene forms of the Indian subcontinent, either ziphodont or not. Contrary to the earlier works on the inter-trappean crocodiles, the present study removes this group as one of the evidences in support of an early (late Cretaceous-early Tertiary) India/Asia collision model. In fact, it provides an additional support for the existence of possible Cretaceous biogeographic links between India, Madagascar, Africa, and South America