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Significance We question a central paradigm: namely, that the protein domain is the “atomic unit” of evolution. In conflict with the current textbook view, our results unequivocally show that duplication of protein segments happens both above and below the domain level among amino acid segments of diverse lengths. Indeed, we show that significant evolutionary information is lost when the protein is approached as a string of domains. Our finer-grained approach reveals a far more complicated picture, where reused segments often intertwine and overlap with each other. Our results are consistent with a recursive model of evolution, in which segments of various lengths, typically smaller than domains, “hop” between environments. The fit segments remain, leaving traces that can still be detected.

Complex evolutionary footprints revealed in an analysis of reused protein segments of diverse lengths