Celestial orientation with the sun not in view: lizards use a time-compensated sky polarization compass

The present investigation was aimed at testing whether the lizard sky polarization compass is time compensated. For this purpose, ruin lizards, Podarcis sicula, were both trained and tested for orientation inside a Morris water maze under clear skies with the sun not in view. During training, lizards showed a striking bimodal orientation along the training axis, demonstrating their capability of determining the symmetry plane of the sky polarization pattern and thus the use of polarization information in orientation. After reaching criteria, lizards were kept 7 days in a 6-h fast clock-shift treatment and then released with the sun not in view. Six-hour clock-shifted lizards showed a bimodal distribution of directional choices, which was oriented perpendicularly to the training axis, as it was expected on the basis of the clock-shift. The results show that the only celestial diurnal compass mechanism that does not need a direct vision of the sun disk (i.e., the sky polarization compass) is a time-compensated compass.