The survival of a bird's egg depends upon its ability to stay within strict thermal limits. 9 Avian eggshell colours have long been considered a phenotype that can help them stay 10 within these thermal limits 1,2 , with dark eggs absorbing heat more rapidly than bright eggs.
11Although disputed 3,4 , evidence suggests that darker eggs do increase in temperature more 12 rapidly than lighter eggs, explaining why dark eggs are often considered as a cost to trade-13 off against crypsis [5][6][7] . Although studies have considered whether eggshell colours can confer 14 an adaptive benefit 4,6 , no study has demonstrated evidence that eggshell colours have 15 actually adapted for this function. This would require data spanning a wide phylogenetic 16 diversity of birds and a global spatial scale. Here we show evidence that darker and 17 browner eggs have indeed evolved in cold climes, and that the thermoregulatory advantage 18 for avian eggs is a stronger selective pressure in cold climates. Temperature alone 19 predicted more than 80% of the global variation in eggshell colour and luminance. These 20 patterns were directly related to avian nesting strategy, such that all relationships were 21 stronger when eggs were exposed to incident solar radiation. Our data provide strong 22 evidence that sunlight and nesting strategies are important selection pressures driving egg 23 pigment evolution through their role in thermoregulation. Moreover, our study advances 24 understanding of how traits have adapted to local temperatures, which is essential if we are 25 to understand how organisms will be impacted by global climate change. 26 27The impact of global climate patterns on the evolution and distribution of traits is an area of 28 increasing importance as global temperatures continue to rise. Birds' eggs are an ideal system for 29 2 exploring the intersection between climate and trait diversity, because a tight thermal range is 30 necessary for the survival of the developing embryo 8 , as eggs are unable to regulate their own 31 temperature 9 . As a result, many birds have adapted incubation behaviours and nest characteristics 32 in response to local conditions 10,11 . In addition to these behavioural adaptations, the adaptive 33 value of eggshell coloration for thermoregulation has been of longstanding interest 1,2,5 . These 34 eggshell colours are generated by just two pigments 12 and eggshell coloration is known to reflect 35 local environmental conditions 13 .
36The white colour found on many eggs (e.g., ostrich eggs) reflects incident solar radiation 37 from their surfaces, but can draw the attention of predators 14 . By contrast, dark brown or heavily 38 speckled eggs (e.g., artic loon eggs) may escape the visual detection of predators, particularly in 39 ground nesting birds 15 , but these darker eggs should heat more rapidly when left in the sun 1,16 . 40 Therefore, in hot climes the thermal costs must be balanced against the adaptive benefits of 41 cryptic pigmentation, while in cold climes thermoregula...