Origin of the Tethyan Hemihoplitidae tested with cladistics (Ancyloceratina, Ammonoidea, Early Cretaceous): an immigration event?

Abstract

The Late Barremian Hemihoplitidae (Ancyloceratina, Ammonoidea) are widely known in the northern Tethyan Margin and the Essaouira-Agadir Basin (Morocco). Their rapid evolution and diversification make them one of the key groups for that period, but their origin remains poorly known and several competing hypotheses have been published. These hypotheses are tested here with cladistic analysis in order to reject those receiving the least support and discuss those well supported. The analysis discards the Crioceratitidae, Emericiceratidae (Emericiceras and Honnoratia) and Toxancyloceras as stem-groups of the Hemihoplitidae (Gassendiceras). The Toxancyloceras appear instead to be a sister-taxon of the Moutoniceras, so we propose the latter to be classified with the Ancyloceratidae rather than with the Heteroceratidae. The best supported hypothesis assumes that the Hemihoplitidae first appeared suddenly in the Essaouira-Agadir Basin at the end of the Early Barremian from small populations of Boreal Paracrioceras. These latter could have migrated southward episodically before invading the northern Tethyan margin at the beginning of the Late Barremian. As a consequence, the Paracrioceratidae fam. nov. is proposed to include the Boreal groups Fissicostaticeras / Paracrioceras / Parancyloceras, and Gassendiceras essaouirae sp. nov. is proposed as a new name for the Moroccan endemic Barrancyloceras maghrebiense sensu Company et al., 2008, non Immel, 1978.