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The main excavations at the Troisième caverne of Goyet in Belgium were conducted by Edouard Dupont in 1868 who identified Palaeolithic human occupations later attributed to the Middle and Upper Palaeolithic. These are represented by an archaeologi- cal record that spans the Mousterian, Lincombian-Ranisian-Jerzmanowician, Aurignacian, Gravettian, and Magdalenian, and then extends into the Neolithic and historic periods. Due to the lack of detailed documentation of the excavated materials, their asso- ciation to a specific chronocultural context has been challenging. Morphometric and taphonomic analyses, combined with direct radiocarbon dating as well as isotopic and genetic analyses, were used to assign human remains to either late Neanderthals or an- cient modern humans from different chronocultural groups. In 2016 the first palaeogenetic investigation of Neanderthal specimens from Goyet was published [1]. Taxonomic assignment was confirmed by performing hybridization capture of the mitochondrial DNA (mtDNA) and later inspecting diagnostic mutations at nucleotide positions that distinguish modern humans from Nean- derthals. Moreover, a phylogenetic reconstruction placed seven nearly complete mtDNA sequences from Goyet within the diver- sity of late Neanderthal mtDNA. An around two-fold coverage nuclear genome was later sequenced from one of those individuals (Goyet Q56-1) [2], revealing a high genetic similarity to other late Neanderthals that is well correlated to their geographical dis- tance. Analyzing modern human remains retrieved at Goyet, mtDNA genomes were initially reported for two specimens directly dated to the Aurignacian, five to the Gravettian, and one to the Magdalenian [3]. Aurignacian-related individuals were particu- larly intriguing as they were found to carry mtDNA haplogroup M, which is almost entirely absent in present-day Europeans. For Gravettian- to Magdalenian-related individuals, the shift from U2/U5 to U8 haplogroups was detected locally - as in other regions of Central Europe - likely influenced by the genetic bottleneck during the Last Glacial Maximum (LGM). Furthermore, nuclear sequences of five modern human individuals from Goyet were produced through genome-wide targeted enrichment [4] revealing local replacement between Aurignacian- and Gravettian-related populations. However, the genetic component associated with a 35,000-year-old individual (Goyet Q116-1) reappeared after the LGM, first in Spain and then in other European regions includ- ing in a Magdalenian-related individual from Goyet (Goyet Q-2). This individual was later found to be the best proxy for a genetic component that was largely displaced in Europe from around 14,000 years ago onwards while surviving in high proportion among Mesolithic individuals from Iberia [5]. Here we present new palaeogenetic data of Neanderthal and modern human individuals from this iconic site. First, we expand the molecular taxonomic identifications with three additional Neanderthal specimens and reconstruct their partial mtDNA genomes. Those confirm the general picture of a limited genetic diversity for late Neanderthals, which is also apparent among the Goyet Neanderthals. Second, working on modern human remains, we produced new mtDNA and nuclear data from four Gravettian specimens. They belong to mtDNA haplogroups U2 and U5, further extending the observa- tion of both mtDNA types being largely present in pre-LGM Europe. Moreover, their nuclear genomes provide additional evidence for the genetic affinity between Gravettian-related groups across Europe, from the present-day regions of the Czech Republic to Belgium and Southern Italy. In conclusion, the deep temporal range covered by the human remains from the Troisième caverne of Goyet provides the unique opportunity to describe within a single archaeological site the major genetic transformations that took place in Europe throughout the Middle and Upper Palaeolithic.

Modern humans have populated Europe for more than 45,000 years1,2. Our knowledge of the genetic relatedness and structure of ancient hunter-gatherers is however limited, owing to the scarceness and poor molecular preservation of human remains from that period3. Here we analyse 356 ancient hunter-gatherer genomes, including new genomic data for 116 individuals from 14 countries in western and central Eurasia, spanning between 35,000 and 5,000 years ago. We identify a genetic ancestry profile in individuals associated with Upper Palaeolithic Gravettian assemblages from western Europe that is distinct from contemporaneous groups related to this archaeological culture in central and southern Europe4, but resembles that of preceding individuals associated with the Aurignacian culture. This ancestry profile survived during the Last Glacial Maximum (25,000 to 19,000 years ago) in human populations from southwestern Europe associated with the Solutrean culture, and with the following Magdalenian culture that re-expanded northeastward after the Last Glacial Maximum. Conversely, we reveal a genetic turnover in southern Europe suggesting a local replacement of human groups around the time of the Last Glacial Maximum, accompanied by a north-to-south dispersal of populations associated with the Epigravettian culture. From at least 14,000 years ago, an ancestry related to this culture spread from the south across the rest of Europe, largely replacing the Magdalenian-associated gene pool. After a period of limited admixture that spanned the beginning of the Mesolithic, we find genetic interactions between western and eastern European hunter-gatherers, who were also characterized by marked differences in phenotypically relevant variants.

This is a reply to the comments of Morey (2014) on our identification of Palaeolithic dogs from several European Palaeolithic sites. In his comments Morey (2014) presents some misrepresentations and misunderstandings that we remedy here. In contrast to what Morey (2014) propounds, our results suggest that the domestication of the wolf was a long process that started early in the Upper Palaeolithic and that since that time two sympatric canid morphotypes can be seen in Eurasian sites: Pleistocene wolves and Palaeolithic dogs. Contrary to Morey (2014), we are convinced that the study of this domestication process should be multidisciplinary.

Numerous bird bones from the Paleocene and early Eocene of the Belgian and Paris basins have been collected by amateur paleontologists. Four bones from the early-middle Selandian of Maret, Belgium are among the earliest Cenozoic avian remains from Europe and include the oldest temporally well constrained records of the Gastornithidae, as well as tentative records of the paleognathous Lithornithidae and the Ralloidea. Another assemblage from the middle Thanetian of Templeuve, France contains multiple bones of the Lithornithidae as well as a record of the Pelagornithidae. Specimens from the latest Thanetian of Rivecourt-Petit Pâtis, France are tentatively assigned to the Ralloidea and Leptosomiformes. An assemblage of 54 bones from the middle Ypresian of Egem, Belgium represents at least 20 species in more than 11 higher-level taxa. Well-identifiable specimens are assigned to the Odontopterygiformes, Galliformes, Messelornithidae, Apodiformes, Halcyornithidae, Leptosomiformes, and Coraciiformes. Further specimens are tentatively referred to the phaethontiform Prophaethontidae and to the Accipitridae, Masillaraptoridae, and Alcediniformes. These three-dimensionally preserved fossils provide new data on the osteology of taxa that are otherwise mainly known from compression fossils with crushed bones. They also further knowledge of the composition of early Paleogene avifaunas of the North Sea Basin. Paleocene avifaunas of Europe and North America appear to have had different compositions and only a few taxa, such as the paleognathous Lithornithidae, are known from both continents. This suggests that the very similar early Eocene avifaunas of Europe and North America are the result of early Cenozoic dispersal events. The well-represented small galliform species from Egem most closely resembles Argillipes aurorum, an ignored galliform species from the London Clay. The tentatively identified fossils of Accipitridae and Alcediniformes would represent the earliest fossil records of these clades. The birds from Egem include few seabirds (Odontopterygiformes, cf. Prophaethontidae) and is dominated by terrestrial species (Galliformes, Messelornithidae). Arboreal birds (Halcyornithidae, Leptosomiformes, cf. Alcediniformes, Coraciiformes) are less abundant and aerial insectivores (Apodiformes) very scarce, which either indicates a taphonomic bias in the composition of the avifauna or particular paleoenvironmental characteristics of the nearshore habitats in that area of the southern North Sea Basin. Grant Information: Funded by Belgian Science Policy Office (project BR/121/A3/PalEurAfrica).