**Coastal Pine-Oak Glacial Refugia in the Mediterranean Basin: A Biogeographic Approach Based on Charcoal Analysis and Spatial Modelling**

**Gaetano Di Pasquale 1, Antonio Saracino <sup>1</sup> , Luciano Bosso 1,\* , Danilo Russo <sup>1</sup> , Adriana Moroni 2, Giuliano Bonanomi <sup>1</sup> and Emilia Allevato 1,\***


Received: 12 May 2020; Accepted: 10 June 2020; Published: 12 June 2020

**Abstract:** During the glacial episodes of the Quaternary, European forests were restricted to small favourable spots, namely refugia, acting as biodiversity reservoirs. the Iberian, Italian and Balkan peninsulas have been considered as the main glacial refugia of trees in Europe. In this study, we estimate the composition of the last glacial forest in a coastal cave of the Cilento area (SW Italy) in seven time frames, spanning from the last Pleniglacial to the Late Glacial. Charcoal analyses were performed in seven archaeological layers. Furthermore, a paleoclimate modelling (Maxent) approach was used to complement the taxonomic identification of charcoal fragments to estimate the past potential distribution of tree species in Europe. Our results showed that the mesothermophilous forest survived in this region in the core of the Mediterranean basin during the Last Glacial Period (LGP, since ~36 ka cal BP), indicating that this area played an important role as a reservoir of woodland biodiversity. Here, *Quercus pubescens* was the most abundant component, followed by a wide variety of deciduous trees and *Pinus nigra*. Charcoal data also pointed at the crucial role of this coastal area, acting as a reservoir for warm temperate trees of genera *Tilia*, *Carpinus* and *Sambucus*, in LGP, in the Mediterranean region. Our modelling results showed that *P. nigra* might be the main candidate as a "*Pinus sylvestris* type" in the study site in the Last Glacial Maximum (LGM). Furthermore, we found that *P. nigra* might coexist with *Q. pubescens* in several European territories both currently and in the LGM. All models showed high levels of predictive performances. Our results highlight the advantage of combining different approaches such as charcoal analysis and ecological niche models to explore biogeographic questions about past and current forest distribution, with important implications to inform today's forest management and conservation.

**Keywords:** Charcoal; Ecological Niche Model; Forest History; Last Glacial Maximum; Maxent; Paleoecology; *Pinus nigra*; *Pinus mugo*/*uncinata*; *Pinus sylvestris*; *Quercus pubescens*
