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Ancient DNA has allowed the study of various aspects of human history in unprecedented detail. However, because the majority of archaic human specimens preserved well enough for genome sequencing have been female, comprehensive studies of Y chromosomes of Denisovans and Neandertals have not yet been possible. Here we present sequences of the first Denisovan Y chromosomes (Denisova 4 and Denisova 8), as well as the Y chromosomes of three late Neandertals (Spy 94a, Mezmaiskaya 2 and El Sidrón 1253). We find that the Denisovan Y chromosomes split around 700 thousand years ago (kya) from a lineage shared by Neandertal and modern human Y chromosomes, which diverged from each other around 370 kya. The phylogenetic relationships of archaic and modern human Y chromosomes therefore differ from population relationships inferred from their autosomal genomes, and mirror the relationships observed on the level of mitochondrial DNA. This provides strong evidence that gene flow from an early lineage related to modern humans resulted in the replacement of both the mitochondrial and Y chromosomal gene pools in late Neandertals. Although unlikely under neutrality, we show that this replacement is plausible if the low effective population size of Neandertals resulted in an increased genetic load in their Y chromosomes and mitochondrial DNA relative to modern humans. ; Q.F. was supported by funding from the Chinese Academy of Sciences (XDB26000000), and the National Natural Science Foundation of China (91731303, 41925009,41630102). A.R. was funded by Spanish government (MICINN/FEDER), grant number CGL2016-75109-P. The reassessment of the Spy collection by H.R., I.C. and P.S. was supported by the Belgian Science Policy Office (BELSPO 2004-2007, MO/36/0112). M.S., M.K. and A.D. were supported by the Russian Foundation for Basic Research (RFBR 17-29-04206). This study was funded by the Max Planck Society and the European Research Council (grant agreement number 694707). ; No

This review summarizes current thinking about the concept of modern behavior in the context of Neandertals and anatomically modern humans. The decoupling of modern anatomy and modern behavior has prompted researchers to reframe studies of the emergence of modern humans as a debate that explicitly focuses on the origins of behavioral modernity making its intersection with modern anatomy a point of discussion rather than a given. Four questions arise from this debate: (a) What is modern behavior? (b) Is the emergence of modern behavior sudden or more gradual? (c) Is modern behavior unique to modern humans or more widely shared with other species, most notably the Neandertals? (d) Is the emergence of modern behavior primarily the result of new cognitive abilities or social, cultural, demographic, and historic factors? This review briefly addresses each of these questions and in the process offers some thoughts on the current state of the debate.