The seven ways eukaryotes produce repeated colour motifs on external tissues

The external tissues of numerous eukaryote species show repeated colour patterns, usually characterized by units that are present at least twice on the body. These dotted, striped or more complex phenotypes carry out crucial biological functions, such as partner recognition, aposematism or camouflage. Very diverse mechanisms explaining the formation of repeated colour patterns in eukaryotes have been identified and described, and it is timely to review this field from an evolutionary and developmental biology perspective. We propose a novel classification consisting of seven families of primary mechanisms: Turing(-like), cellular automaton, multi-induction, physical cracking, random, neuromuscular and printing. In addition, we report six pattern modifiers, acting synergistically with these primary mechanisms to enhance the spectrum of repeated colour patterns. We discuss the limitations of our classification in light of currently unexplored extant diversity. As repeated colour patterns require both the production of a repetitive structure and colouration, we also discuss the nature of the links between these two processes. A more complete understanding of the formation of repeated colour patterns in eukaryotes will require (i) a deeper exploration of biological diversity, tackling the issue of pattern elaboration during the development of non-model taxa, and (ii) exploring some of the most promising ways to discover new families of mechanisms. Good starting points include evaluating the role of mechanisms known to produce non-repeated colour patterns and that of mechanisms responsible for repeated spatial patterns lacking colouration.