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B.W. was supported by the NERC and the Leverholme Trust and is now supported by the Henry Luce Foundation; M.C. is supported by the Wellcome Trust. We are grateful to A. Cain for the translation of Linnaeus' Systema Naturae (10th edition, 1758); A. Chamberlain for making his data available; and A. Chamberlain, W. Kimbel, D. Lieberman, and D. Strait for commenting on the manuscript.
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