1 1 Unique evolution of vitamin A as an external pigment in 2 tropical starlings 3 4 Ismael Galván1*, Khaled Murtada2,3, Alberto Jorge4, Ángel Ríos2,3, and 5 Mohammed Zougagh2,5 6 7 1Department of Evolutionary Ecology, Doñana Biological Station, Consejo Superior de 8 Investigaciones Científicas (CSIC), 41092 Sevilla, Spain 9 2Regional Institute for Applied Scientific Research (IRICA), University of Castilla-La 10 Mancha, 13071 Ciudad Real, Spain 11 3Department of Analytical Chemistry and Food Technology, Faculty of Chemical Science 12 and Technology, University of Castilla-La Mancha, 13071 Ciudad Real, Spain 13 4Laboratory of Non-Invasive Analytical Techniques, National Museum of Natural Sciences, 14 Consejo Superior de Investigaciones Científicas (CSIC), 28006 Madrid, Spain 15 5Department of Analytical Chemistry and Food Technology, Faculty of Pharmacy, 16 University of Castilla-La Mancha, 02071 Albacete, Spain 17 18 *Author for correspondence (
[email protected]) 19 20 Running title: Evolution of retinol-based pigmentation 21 22 KEY WORDS: Biological pigments, Birds, Carotenoids, Color evolution, Retinol 23 24 25 2 26 Summary statement 27 We have discovered a new class of external biological pigments, after showing that an 28 Afrotropical starling has physiological capacity to deposit retinol in its integument and 29 create bright yellow coloration. 30 31 ABSTRACT 32 Pigments are largely responsible for the appearance of organisms. Most biological 33 pigments derive from the metabolism of shikimic acid (melanins), mevalonic acid 34 (carotenoids) or levulinic acid (porphyrins), which thus generate the observed diversity of 35 external phenotypes. Starlings are generally dark birds despite iridescence in feathers, but 36 10 % of species have evolved plumage pigmentation comprising bright colors that are 37 known to be produced only by carotenoids.