Anurans from the Candeleros Formation (?Cenomanian-Turonian) of west-central Argentina: new evidence for pipoid evolution

The fluvio-lacustrine deposits of the ?Cenomanian-Turonian Candeleros Formation in the northwestern fringe of Patagonia have yielded numerous remains of vertebrates, including anurans. A new, partially articulated immature specimen of a small pipimorph pipoid from the upper part of this unit is described herein. Although incompletely preserved, the morphology and proportions of the skeleton are strongly reminiscent of corresponding elements of the holotype of Avitabatrachus uliana from the lower section of the same formation at a nearby locality, thus suggesting that these specimens might represent the same, or a closely related, taxon. The new specimen is a metamorphosing individual: it has a zygapophyseal articulation between the sacrum and the neural arch of a postsacral vertebra that bears conspicuous transverse processes, and four pairs of ribs. These traits are not present in the available material of A. uliana, but they might have occurred within the normal range of variation of a single species. Although the generic and specific allocation of the new material is uncertain, a parsimony analysis performed to clarify the relationships of A. uliana supports a phylogenetic placement of this taxon outside the crown Pipidae. The evidence provided by the Candeleros anurans confirms the presence of stem pipids in Patagonia in the early Late Cretaceous.