What, if anything, is the adaptive function of countershading?

Countershading, the gradation of colour from dark on the dorsum to light on the ventrum, is generally considered to have the effect of making organisms difficult to detect. The mechanism that facilitates this form of crypsis is often considered to be concealment of shadows cast on the body of the animal. We review the current empirical evidence for the cryptic function of countershading and for the mechanism underlying it. We argue that there is no conclusive evidence that countershading per se provides any selective advantage in terrestrial or aerial environments. However, the highly refined adaptations of some marine organisms to match the different background light conditions against which they are set when viewed from different aspects strongly suggest an adaptive advantage to countershading in these environments. In none of the cases discussed in this review was the conventional explanation of self-shadow concealment a more plausible explanation for countershading than the alternative explanation that the dorsum and ventrum experience different selection pressures (often associated with background matching).