Distinctive colour patterns in dogs are an integral component of canine diversity. Colour pattern differences are thought to have arisen from mutation and artificial selection during and after domestication from wolves but important gaps remain in understanding how these patterns evolved and are genetically controlled. In other mammals, variation at the ASIP gene controls both the temporal and spatial distribution of yellow and black pigments. Here, we identify independent regulatory modules for ventral and hair cycle ASIP expression, and we characterize their action and evolutionary origin. Structural variants define multiple alleles for each regulatory module and are combined in different ways to explain five distinctive dog colour patterns. Phylogenetic analysis reveals that the haplotype combination for one of these patterns is shared with Arctic white wolves and that its hair cycle-specific module probably originated from an extinct canid that diverged from grey wolves more than 2 million years ago. Natural selection for a lighter coat during the Pleistocene provided the genetic framework for widespread colour variation in dogs and wolves.

狗独特的颜色模式是犬类多样性的一个组成部分。颜色模式差异被认为是由狼驯化期间和之后的突变和人工选择引起的，但在理解这些模式如何进化和遗传控制方面仍然存在重要差距。在其他哺乳动物中，ASIP基因的变异控制着黄色和黑色色素的时间和空间分布。在这里，我们确定了腹侧和毛发周期ASIP的独立监管模块表达，我们描述了它们的行为和进化起源。结构变异为每个调节模块定义了多个等位基因，并以不同的方式组合以解释五种不同的狗颜色模式。系统发育分析表明，其中一种模式的单倍型组合与北极白狼共享，并且其毛发周期特异性模块可能起源于 200 万多年前从灰狼中分化出来的已灭绝犬科动物。更新世期间对浅色皮毛的自然选择为狗和狼的广泛颜色变异提供了遗传框架。