Aim The aims of this study were to assess the distribution of putative Mediterranean refugia of plants, to compare the locations of refugia and those of regional hotspots of plant biodiversity, and to provide a critical analysis of the Mediterranean refugium paradigm. Furthermore, we consider how biogeographical and genetic results can be combined to guide global conservation strategies.Location The Mediterranean region.Methods We started from a detailed analysis of the scientific literature (1993)(1994)(1995)(1996)(1997)(1998)(1999)(2000)(2001)(2002)(2003)(2004)(2005)(2006)(2007) in order to identify refugia in the Mediterranean region, based on intraspecific phylogeographical studies of plant species. We used population locations together with gene-pool identity to establish the database, comparing patterns of phylogeographical concordance with the locations of Mediterranean refugia. We then tested the biogeographical congruence between two biodiversity components, namely phylogeographical refugia and regional hotspots. ResultsWe identified 52 refugia in the Mediterranean bioclimatic region and confirmed the role played by the three major peninsulas, with a shared total of 25 refugia. We emphasize the importance of areas that have previously been attributed a lesser role (large Mediterranean islands, North Africa, Turkey, Catalonia). Of the 52 refugia identified, 33 are situated in the western Mediterranean Basin and 19 in the eastern part. The locations of the phylogeographically defined refugia are significantly associated with the 10 regional hotspots of plant biodiversity, with 26 of these refugia (i.e. 50%) occurring within the hotspots.Main conclusions The locations of refugia are determined by complex historical and environmental factors, the cumulative effects of which need to be considered because they have occurred since the Tertiary, rather than solely during the last glacial period. Refugia represent climatically stable areas and constitute a high conservation priority as key areas for the long-term persistence of species and genetic diversity, especially given the threat posed by the extensive environmental change processes operating in the Mediterranean region. The refugia defined here represent 'phylogeographical hotspots'; that is, significant reservoirs of unique genetic diversity favourable to the evolutionary processes of Mediterranean plant species.

Aim Phylogeography of fruit trees is challenging due to recurrent exchanges between domesticated and wild populations. Here we tested the eastern refugium hypothesis (ERH) for the carob tree, Ceratonia siliqua, which supports its natural and domestication origins in the eastern Mediterranean and a feral origin in the west. Location Mediterranean basin. Taxon Ceratonia siliqua L., Leguminosae. Methods A phylogenetic reconstruction based on two nuclear and one plastid sequences was performed to estimate the divergence time between the carob tree and its sister species, Ceratonia oreothauma. Variation from four plastid regions and 17 nuclear microsatellite loci were used to decipher genetic structure in the carob tree and to test coalescent‐based models by an Approximate Bayesian computation (ABC) approach. We assessed our hypotheses by examining palaeobotanical records and hindcasting the past distribution of the carob tree at Mid‐Holocene, Last Glacial Maximum (LGM) and Last Interglacial (LIG) using species distribution modelling. Results The split between C. oreothauma and C. siliqua was estimated at 6.4 Ma, and a first divergence within C. siliqua at 1.3 Ma. After a continuous presence since the Oligocene, Ceratonia was rarely found in the fossil record during the Pleistocene but present in the western and the eastern Mediterranean. Plastid and nuclear markers, characterized by low allelic richness, revealed a strong west‐east genetic structuring. ABC analyses rejected the ERH. Main conclusions Our study supports a severe population decline during LIG. The strong west–east divergence and the occurrence of four lineages within C. siliqua provided support for a new hypothesis of multiple domestications of the carob tree from native populations throughout the Mediterranean basin.

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