Southern dispersal and Palaeoecological implications of woolly rhinoceros ( Coelodonta antiquitatis): review of the Iberian occurrences

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Cold-adapted large mammal populations spread southward during the coldest and driest phases of the Late Pleistocene reaching the Iberian Peninsula. Presence of woolly rhinoceros ( Coelodonta antiquitatis) can be identified from 23 Iberian sites, which is compiled and analyzed herein, and the fossil specimens from seven of these sites are described here for first time. Morphological and biometrical analyses demonstrate that the Iberian woolly rhinoceros did not significantly differ from individuals of other European populations, but represent the westernmost part of a continuous Eurasian belt of distribution. The first presence of woolly rhino in the Iberian Peninsula has been identified during the late Middle Pleistocene and early Late Pleistocene. However, the highest abundance of this species is recorded during MIS 3 and 2. The latest Iberian occurrences can be dated around 20 ka BP. The presence of woolly rhinoceros in the Iberian Peninsula correlates with periods of extreme dry and cold climatic conditions documented in Iberian terrestrial and marine sediment sequences. From a palaeobiogeographic point of view, the maximum southern spread of C. antiquitatis on the Iberian Peninsula was registered during the late Middle Pleistocene or early Late Pleistocene, reaching the latitude of Madrid (about 40°N). Subsequently, during MIS 3 and 2, all Iberian finds were restricted to the Northern regions of Iberia (Cantabrian area and Catalonia). The southern expansion of C. antiquitatis during the Late Pleistocene in the Iberian Peninsula reached similar latitudes to other Eurasian regions. The ecological composition of fossil assemblages with presence of woolly rhinoceros was statistically analyzed. Results show that temperate ungulate species are predominant at Iberian assemblages, resulting in a particular mixture of temperate and cold elements different of the typical Eurasian cold-adapted faunal associations. This particular situation suggests two possible explanations: a) Eventual migrations during the coldest time spans, resulting in a mixing of cold and temperate faunas, instead a faunal replacing; b) Persistence of woolly rhinoceros populations in the Iberian Peninsula during interglacial episodes confined at cryptic southern refugia.