Tracking the last elephants in Europe during the Würm Pleniglacial: the importance of the Late Pleistocene aeolianite record in SW Iberia

Abstract In his short joint ventures across the world of vertebrate tracks, Richard Bromley recognized the aeolian sands as unsuitable soft substrates for their preservation. Only after his work in the Balearic Islands, a more systematic study of coastal aeolianites worldwide revealed that these depositional systems could preserve a highly important record of behavioural trace fossils for the evolution of vertebrates, especially in the Pleistocene, including extinct megafauna and the escalation of the Homo. Here we describe coastal aeolianites from the upper Pleistocene of SW Iberia, namely SW Portugal and Gibraltar, with trackways, tracks and trampled surfaces of the last elephants in mainland Europe. Photogrammetric 3 D modelling and analysis of the relevant proboscidean track levels allowed revision of and support for previous ichnotaxonomic identification to Proboscipeda panfamilia and behavioural interpretations of the producer. Smaller and very large trackways and footprints attributed to Palaeoloxodon antiquus are described and discussed according to new and recent, but always rare findings. The seeming progressive and definitive extinction of this species towards southern Iberia, following the same pattern for the replacement of the Neanderthals during the last initial Pleniglacial (until ca. 28 ka), suggests evidence for co-evolution.