Solving the missing pieces of the gharial puzzle: new phylogenetic framework combining morphological, molecular, and biostratigraphic data to unravel the evolution of long-snouted crocodylians.

Among the extant crocodylians are two species with long, narrow snouts: Gavialis gangeticus,the Indian gharial and Tomistoma schlegelii, the "false" gharial. These enigmatic species are considered by the IUCN red list as critically endangered and vulnerable, respectively. However, despite this, knowledge of their evolutionary history is lacking. Extensive debate has surrounded the gharials for over four decades and remains unsolved today: the so-called gharial problem. Whereas molecular studies consistently indicate that these two species are sister taxa, morphological studies of both living and fossil taxa find that they belong to distantly related lineages. Moreover, molecular clock estimates indicate a shallow divergence time of 18-31 million years ago. This entirely contradicts the rich fossil record of gharials: in contrast to the modern gharials, these fossil taxa comprise a huge diversity and suggest that tomistomines and gavialines have diverged from each other at least 70 million years ago, prior to the K/Pg mass extinction.

European museums, and especially the Royal Belgian Institute of Natural Sciences, Brussels, comprise rich collections containing many of the oldest fossil gavialoids, crucial to solving the gharial problem. Nevertheless, few modern morphological studies have been performed on these specimens, and their stratigraphic age is often poorly constrained. Therefore, in a new project we will use a multidisciplinary approach to study these specimens, combining morphological study and biostratigraphic analyses using dinoflagellate cysts. Moreover, we will revise the classical methods used by paleontologists to study fossil crocodylians, devising a new phylogenetic framework that makes use of both morphological, molecular, and biostratigraphic data. Here, we will present some of the first preliminary results of this project.