



Boxes and whiskers represent 25% quartile, 75% quartile, minimum, and maximum, with horizontal line at median. S2) were highly correlated with measured size in 375 dogs (Pearson correlation). ( D) Owner responses to the size question (fig. ( B and C) Surveys include (B) 79 published behavior questions ( 34– 37) and 39 new questions about heritable, easy-to-identify traits that fall into (C) four categories that potentially vary in heritability, including physical traits (fig. ( A) Selection on behavior in dogs predates modern breeds, which have existed for ~1% of dog history ( 10). The extent to which ancient behavioral propensities persist in modern breeds, defined primarily by aesthetics and often disconnected from functional behavioral selection, is unclear. Modern breeds retain these component motor patterns, but their contexts, sequences, and thresholds vary ( 12, 13). Before the 1800s, dogs were primarily selected for functional roles such as hunting, guarding, and herding ( 11)-heritable behaviors derived from the wolf predatory sequence ( 12). By contrast, the modern dog breed, emphasizing conformation to a physical ideal and purity of lineage, is a Victorian invention ( 10). Early humans may have given favored dogs increased access to limited resources, but there is little evidence of humans intentionally breeding dogs until 2000 years ago ( 8, 9). Prehistoric wolves likely adapted to use human refuse through changes in morphology, behavior, metabolism, and reproduction ( 3– 7).

Modern dog breeds are less than 160 years old (~50 to 80 generations), a blink in evolutionary history compared with the origin of dogs more than 10,000 years ago ( 1, 2) ( Fig. Karlsson +22 authors +20 authors +15 authors fewer Authors Info & Affiliations White, Zhiping Weng, Andrés Colubri, Diane P. Kathleen Morrill, Jessica Hekman, , Xue Li, Jesse McClure, , Brittney Logan, Linda Goodman, Mingshi Gao, Yinan Dong, Marjie Alonso, , Elena Carmichael, Noah Snyder-Mackler, Jacob Alonso, Hyun Ji Noh, Jeremy Johnson, Michele Koltookian, Charlie Lieu, Kate Megquier, Ross Swofford, Jason Turner-Maier, Michelle E.
