The youngest pangolin (Mammalia, Pholidota) from Europe

The fossil record for pangolins is sparse. Current biogeographic data suggest this clade originated in Europe, though pangolins seem to have disappeared from the European paleontological record during the middle Miocene, when they were hypothesized to have been pushed toward more tropical and sub-tropical equatorial environments due to global cooling trends. Here we report on a nearly complete humerus of a pangolin from the early Pleistocene (similar to 2.2-1.9 Ma) site of Graunceanu, Romania. This fossil revises this previous understanding of pangolin evolution and biogeography and represents both the youngest fossil pangolin from Europe and the only fossil from Pleistocene Europe. The new species described here, Smutsia olteniensis, sp. nov., shares several synapomorphic traits with other Smutsia species, which are currently found only in Africa. However, relative to extant Smutsia, it has several unique traits that set it apart, including a longer and narrower entepicondyle, an enlarged supinator crest, and an enlarged greater tubercle. Together this unique suite of features justifies the description of a new species. This specimen definitively demonstrates that pangolins were present in Europe during the Pleistocene. Further, Smutsia has previously been thought to be an African taxon, with the oldest specimen from South Africa at similar to 5 Ma and living species found across Africa. This specimen now demonstrates that Smutsia previously had a far larger biogeographic range. Finally, Graunceanu has been reconstructed to have consisted of relatively open grasslands and woodlands, which is an unusual habitat for most pangolins.