Revision of Vaccinites loftusi (Woodward, 1855) (Bivalvia: Hippuritida) and proposal of three new Vaccinites species from the Campanian–Maastrichtian of Eurasia

The revision of Vaccinites loftusi (Woodward) comprised the critical analysis of issued descriptions and figures, together with the study of the type specimens and newly collected material from the type locality in Turkey, and also from western Bulgaria, eastern Serbia, and Spain. Five (three of them new) different although phylogenetically related species are characterized, each one arisen in a different area of the Mediterranean Tethys. Main diagnostic characters considered are size and ontogenetic development of the ligament ridge and secondary pillars, while angular measures of pillar–myo-cardinal arrangement and main infolds setting, although clustered for each species, depict extensive overlap. Vaccinites alceotarlaoi sp. nov., from the lower Campanian of Serbia and Spain, appears as the most primitive and the origin of the other species. Vaccinites pirenaicus sp. nov., Vaccinites bilottei sp. nov., and Vaccinites loftusi arose at the late Campanian in Pyrenees, Bulgaria, and Middle East, respectively. Vaccinites syriaca Vautrin originated from V. loftusi at the early Maastrichtian. The purported origin of the Eurasian multiple-fold hippuritids (Pironaea spp) from any of these species, with short secondary pillars not affecting the radial pattern of the canal system of the left valve, is rejected.