A revision of the Lari (Aves, Charadriiformes) from the early Miocene of Saint-Gérand-le-Puy (Allier, France)

Three species of gull-like birds were described by Milne-Edwards in the 19th century from the early Miocene of Saint-Gérand-le-Puy, France, two of which have been since then moved to the genus Laricola. These fossils are redescribed and revised in the present study. We further describe two new species of the taxon Laricola, L. intermedia, sp. nov., and L. robusta, sp. nov., as well as two species of a new taxon Sternalara, S. minuta and S. milneedwardsi, thus substantially increasing the number of charadriiform taxa known from this locality. To find out how Laricola relates to modern taxa, we performed a phylogenetic analysis of 41 osteological characters, which resulted in Laricola being placed outside of Lariidae, as early representatives of the newly designated group Laromorphae, which encompasses terns, gulls, and skimmers. The results of our analysis have additionally enabled us to evaluate relationships between extant taxa of the Laromorphae.