Introduction
Dogs (Canis lupus familiaris), the first species to be domesticated by humans, were trained not only for practical roles such as guarding, assisting in hunting, and providing companionship, but they also held significant importance as offerings in funerary rituals and other ceremonial practices. This underscores their integral role in human cultural expressions. In China, the tradition of burying dogs alongside humans can be traced back to the Jiahu archaeological site in Wuyang, dating back to the Neolithic period (~8500BP) (Henan Provincial Institute of Cultural Heritage and Archaeology, 1999). During the Shang Dynasty, the ritual of dog sacrifice underwent a series of evolutionary stages, with its prevalence notably increasing in the late Shang period (~3000BP).The most representative sites are the Yinxu site in Anyang, Henan Province (The Anyang Archaeological Team, 1979), the Taixi site in Gaocheng, Hebei Province (Hebei Provincial Institute of Cultural Relics and Archaeology, 1985) and the Zhoujiazhuang site in this study. Previous studies (Dong, 2020; Guo, 2014; Liu, 2011, 2016) have explored aspects such as the sacrificial position of dogs, their functions, and the human–dog relationship. However, the genetic origins, domestication,and migratory processes of these dogs remain unclear.Current research on ancient DNA (aDNA) concerning dogs primarily focuses on the origins and migration patterns of domestic dogs. Mitochondrial DNA (mtDNA) studies have been particularly instrumental in this discourse, suggesting that dogs may have originated in several regions, including Europe (Thalmann et al., 2013), East Eurasia (Zhang et al., 2024), Siberia (Da Silva Coelho et al., 2021; Perri et al., 2021), and Southern East Asia (Pang et al., 2009; Wang et al., 2016; Zhang et al.,2020). In addition to these findings, research on nuclear genomes and Ychromosomes has identified two further centres of domestication in Central Asia and the Middle East (Gray et al., 2010; Shannon et al., 2015;Vonholdt et al., 2010) and has proposed the possibility of independent domestication events occurring in the eastern and western parts of Eurasia (Bergstrom ¨ et al., 2022; Frantz et al., 2016).Moreover, studies have revealed genetic variations and migration patterns of domestic dogs across diverse geographical regions and historical periods (Ameen et al., 2019; Bergstrom ¨ et al., 2020; Feuerborn et al., 2021; Greig et al., 2018; Ollivier et al., 2018; Zhang et al., 2020,2024), providing essential insights for a deeper understanding of the origins of domestic dogs and the intricate relationship between humans and canines. However, the population composition, origin, and migration pathways of domestic dogs in East Asia remain unclear, particularly during the Late Shang period, and the specific relationship between the Shang people and dogs has not been adequately elucidated.
