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This is actually completely backwards. Unborn animals always start out with no color (white) and the color spreads from the spine outwards. We should never think of white as a marking covering up color. Instead white areas are a failure in normal development.

Oh wait they do have it right cause it states "pigment migrates to different parts of the body during the development of the embryo" in the article. However they also refer to white as "creeping" and "spreading" which would suggest that the embryo is colored to start so I could see how it would be confusing.

Yea they have the link for that site in the article. I was reading through it and its very interesting. I have three tricolor terriers and its neat to see how two black & tan coloring similar to a doberman and the other has the saddle/creeping tan.