The Viking maritime expansion from Scandinavia (Denmark, Norway, and Sweden) marks one of the swiftest and most far-flung cultural transformations in global history. During this time (c. 750 to 1050 CE), Viking sailors reached Greenland, North America, and large parts of western Eurasia, and left a legacy that persists today. To understand the genetic structure and influence of the Viking expansion, we sequenced the genomes of 442 ancient humans from across Europe and Greenland ranging from the Bronze Age (c. 2400 BCE) to the early Modern period (c. 1600 CE), with particular emphasis on the Viking Age. We find that the period preceding the Viking Age was accompanied by foreign gene flow into Scandinavia from the south and east: spreading from Denmark and eastern Sweden to the rest of Scandinavia. Despite the close linguistic similarities of modern Scandinavian languages, we observe genetic structure within Scandinavia, suggesting that regional population differences were already present 1,000 years ago. We find evidence for a majority of Danish Viking presence in England, Swedish Viking presence in the Baltic, and Norwegian Viking presence in Ireland, Iceland, and Greenland. Additionally, we see substantial foreign European ancestry entering Scandinavia during the Viking Age. We also find that several of the members of the only archaeologically well-attested Viking expedition were close family members. By comparing Viking Scandinavian genomes with present-day Scandinavian genomes, we find that pigmentation-associated loci have undergone strong population differentiation during the last millennia. Finally, we are able to trace the allele frequency dynamics of positively selected loci with unprecedented detail, including the lactase persistence allele and various alleles associated with the immune response. We conclude that the Viking diaspora was characterized by substantial foreign engagement: distinct Viking populations influenced the genomic makeup of different regions of Europe, while Scandinavia also experienced increased contact with the rest of the continent.

In this study we explored quantitatively the relationships between the size of bats, the frequencies in their echolocation calls, and the incidence of moths and beetles in their diets. We focused on the predictions of the allotonic frequency hypothesis which states that some insectivorous bats increase their access to moths that can hear echolocation calls by shifting to frequencies to which the ears of these insects are less sensitive. The hypothesis predicts that the frequencies dominating the echolocation calls of bats may be correlated with the incidence of moths in their diets. We collected data for 62 species of bats that take airborne prey, usually¯ying insects, 25 species of high duty cycle echolocating bats (Rhinolophidae and Hipposideridae) and 37 species that are low duty cycle echolocators (Vespertilionidae and Molossidae). For bats whose echolocation calls are dominated by frequencies < 100 kHz our regression analyses showed a parabolic dependency between moth consumption (% volume or % frequency) and echolocation call frequency (kHz), supporting the allotonic frequency hypothesis. The use of echolocation calls dominated by frequencies outside the range of best hearing by moths may indeed increase the availability of these insects to the bats. However, when the same analysis was performed with only the bat species using echolocation calls dominated by sounds > 100 kHz, the relationship was not statistically signi®cant, suggesting that morphological characteristics rather than echolocation call frequency may limit the range of potential prey items. Our analyses also demonstrate the importance of jaw morphology as a predictor of the incidence of beetles or moths in the diets of bats, and reveal that generally bigger species (as de®ned by forearm length) use echolocation calls dominated by lower frequencies than smaller species. In both high duty and low duty cycle echolocating bats the relationship between body size and dominant call frequency was best described by a linear model. We also propose that perch hunting was central in the development of the high duty cycle approach to echolocation.

87 The Viking maritime expansion from Scandinavia (Denmark, Norway, and Sweden) marks one 88 of the swiftest and most far-flung cultural transformations in global history. During this time 89 (c. 750 to 1050 CE), the Vikings reached most of western Eurasia, Greenland, and North 90 America, and left a cultural legacy that persists till today. To understand the genetic structure 91 and influence of the Viking expansion, we sequenced the genomes of 442 ancient humans from 92 across Europe and Greenland ranging from the Bronze Age (c. 2400 BC) to the early Modern 93 period (c. 1600 CE), with particular emphasis on the Viking Age. We find that the period 94 preceding the Viking Age was accompanied by foreign gene flow into Scandinavia from the 95 south and east: spreading from Denmark and eastern Sweden to the rest of Scandinavia. 96Despite the close linguistic similarities of modern Scandinavian languages, we observe genetic 97 structure within Scandinavia, suggesting that regional population differences were already 98 present 1,000 years ago. We find evidence for a majority of Danish Viking presence in England, 99 Swedish Viking presence in the Baltic, and Norwegian Viking presence in Ireland, Iceland, and 100Greenland. Additionally, we see substantial foreign European ancestry entering Scandinavia 101 during the Viking Age. We also find that several of the members of the only archaeologically 102 well-attested Viking expedition were close family members. By comparing Viking Scandinavian 103 genomes with present-day Scandinavian genomes, we find that pigmentation-associated loci 104 have undergone strong population differentiation during the last millennia. Finally, we are able 105 to trace the allele frequency dynamics of positively selected loci with unprecedented detail, 106 including the lactase persistence allele and various alleles associated with the immune response. 107We conclude that the Viking diaspora was characterized by substantial foreign engagement: 108 distinct Viking populations influenced the genomic makeup of different regions of Europe, 109 while Scandinavia also experienced increased contact with the rest of the continent. 110 111