Fig. 2.

Multimodal compasses in insects. a Insects derive their orientation with respect to the solar and lunar azimuths either through direct observation (1.) or indirect measures (2. detection of the contrasolar/contralunar azimuth which has the highest degree of polarisation; 3. measurement of chromatic gradients; 4. sampling the polarised light pattern). b Insects also orient with respect to prominent visual features such as the visual panorama (top) or the milky way (bottom). Background fisheye images are sampled from dung beetle habitats in South Africa and provided by Dr James Foster. c Consistent wind provides a short-term orientation cue known to be used by ants and dung beetles. d Insects derive their orientation with respect to the Earth’s North–South axis but the sensory pathways remain unresolved. e Proprioception tracks changes in animal heading iteratively, which is suitable for direction tracking over short durations, for instance, changing direction alters haltere orbits. Insect species combine these multimodal and multiscale compasses differently depending on their navigational need, for example, simple course stabilisation in dung-beetles, central-place navigation in ants, and long-range migration in butterflies