We assembled genome-wide data from 271 ancient Iberians, of whom 176 are from the largely unsampled period after 2000 BCE, thereby providing a high-resolution time transect of the Iberian Peninsula. We document high genetic substructure between northwestern and southeastern hunter-gatherers before the spread of farming. We reveal sporadic contacts between Iberia and North Africa by~2500 BCE and, by~2000 BCE, the replacement of 40% of Iberia's ancestry and nearly 100% of its Y-chromosomes by people with Steppe ancestry. We show that, in the Iron Age, Steppe ancestry had spread not only into Indo-European-speaking regions but also into non-Indo-European-speaking ones, and we reveal that present-day Basques are best described as a typical Iron Age population without the admixture events that later affected the rest of Iberia. Additionally, we document how, beginning at least in the Roman period, the ancestry of the peninsula was transformed by gene flow from North Africa and the eastern Mediterranean.

The spread of farming out of the Balkans and into the rest of Europe followed two distinct routes: An initial expansion represented by the Impressa and Cardial traditions, which followed the Northern Mediterranean coastline; and another expansion represented by the LBK (Linearbandkeramik) tradition, which followed the Danube River into Central Europe. Although genomic data now exist from samples representing the second migration, such data have yet to be successfully generated from the initial Mediterranean migration. To address this, we generated the complete genome of a 7,400-year-old Cardial individual (CB13) from Cova Bonica in Vallirana (Barcelona), as well as partial nuclear data from five others excavated from different sites in Spain and Portugal. CB13 clusters with all previously sequenced early European farmers and modern-day Sardinians. Furthermore, our analyses suggest that both Cardial and LBK peoples derived from a common ancient population located in or around the Balkan Peninsula. The Iberian Cardial genome also carries a discernible hunter–gatherer genetic signature that likely was not acquired by admixture with local Iberian foragers. Our results indicate that retrieving ancient genomes from similarly warm Mediterranean environments such as the Near East is technically feasible.

Marine food–reliant subsistence systems such as those in the African Middle Stone Age (MSA) were not thought to exist in Europe until the much later Mesolithic. Whether this apparent lag reflects taphonomic biases or behavioral distinctions between archaic and modern humans remains much debated. Figueira Brava cave, in the Arrábida range (Portugal), provides an exceptionally well preserved record of Neandertal coastal resource exploitation on a comparable scale to the MSA and dated to ~86 to 106 thousand years ago. The breadth of the subsistence base—pine nuts, marine invertebrates, fish, marine birds and mammals, tortoises, waterfowl, and hoofed game—exceeds that of regional early Holocene sites. Fisher-hunter-gatherer economies are not the preserve of anatomically modern people; by the Last Interglacial, they were in place across the Old World in the appropriate settings.

The Middle Pleistocene is a crucial time period for studying human evolution in Europe, because it marks the appearance of both fossil hominins ancestral to the later Neandertals and the Acheulean technology. Nevertheless, European sites containing well-dated human remains associated with an Acheulean toolkit remain scarce. The earliest European hominin crania associated with Acheulean handaxes are at the sites of Arago, Atapuerca Sima de los Huesos (SH), and Swanscombe, dating to 400–500 ka (Marine Isotope Stage 11–12). The Atapuerca (SH) fossils and the Swanscombe cranium belong to the Neandertal clade, whereas the Arago hominins have been attributed to an incipient stage of Neandertal evolution, to Homo heidelbergensis, or to a subspecies of Homo erectus. A recently discovered cranium (Aroeira 3) from the Gruta da Aroeira (Almonda karst system, Portugal) dating to 390–436 ka provides important evidence on the earliest European Acheulean-bearing hominins. This cranium is represented by most of the right half of a calvarium (with the exception of the missing occipital bone) and a fragmentary right maxilla preserving part of the nasal floor and two fragmentary molars. The combination of traits in the Aroeira 3 cranium augments the previously documented diversity in the European Middle Pleistocene fossil record

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Coastal areas as reservoirs of resources for hominid groups have been widely studied in recent years. These areas combine marine with terrestrial and wetland resources and would have been optimum sites for hominids, including Neanderthals. This is the case with the Cova del Gegant, a cave that today opens directly onto the Mediterranean Sea and is located in the north-eastern Iberian Peninsula. The geomorphological evolution of the Massis del Garraf has provided evidence that during the late Pleistocene there was a littoral platform between 8 and 13 km wide in front of the Cova del Gegant. Within this framework, the data derived from analysis of the small vertebrates and large mammals recovered from Cova del Gegant, including taxa currently absent from the Massis del Garraf, suggest that the landscape surrounding the cave provided a richer terrestrial ecosystem for Neanderthals than is available in this zone today. Analysis of the small-vertebrate association from the cave reveals that the landscape surrounding the cave was dominated by woodland-edge and open environments and that the climate was Mediterranean. The results have been compared with the only Iberian site with similar characteristics to the Cova del Gegant, Gorham's cave (southern Iberia, Gibraltar), revealing differences and similarities in the landscape and climate on the basis of the small-mammal assemblages as well as the differences in the accessibility to terrestrial mammalian resources for the Neanderthal groups. The landscape and the climate were reasonably similar at the two sites, but the differences in the accessibility of resources for the Neanderthals are directly related to the location of the sites and the coastal position. Whereas the Cova del Gegant was on a route of mammal migration (between the Ebro Valley and France) suited for securing terrestrial resources, Gorham's cave is located on a small peninsula with a lower abundance of terrestrial mammal resources. This is probably why the Neanderthal groups at Gorham's cave exploited marine resources, whereas there is no evidence of marine resources having been exploited at Cova del Gegant, even though the seashore was nearby.