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Eugene A. Carver, Coat color genetics of the German shepherd dog, Journal of Heredity, Volume 75, Issue 4, July 1984, Pages 247–252, https://doi.org/10.1093/oxfordjournals.jhered.a109926
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Abstract
The coat-color genetics of the German shepherd dog conforms in most cases to the scheme proposed by Little. In the agouti series, the alleles ay, aw and at are present in the order proposed. However, the dominant black allele AS is absent and the black dogs in this breed are due to a recessive agouti black allele, a. In the extension series, there is evidence for the two alleles E and e as proposed by Little but insufficient evidence to confirm or deny the existence of the other alleles Em and ebr. In the albino series, the alleles C and cch are apparently present and behave as originally suggested. Other phaeomelanin modifiers that are present include the lljin int series and possibly polygenic modifiers. Black-and-tan dogs with patches of gray hairs invading the tan areas and some of the black areas are due to a recessive gene at a new locus. White dogs also are due to a single-factor recessive w that produces a white coat while not affecting the eye or skin pigments.