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Monarch butterflies (Danaus plexippus) breeding in eastern North America are famous for their annual fall migration to their overwintering grounds in Mexico. However, the mechanisms they use to successfully reach these sites remain poorly understood. Here, we test whether monarchs are true navigators who can determine their location relative to their final destination using both a "compass" and a "map". Using flight simulators, we recorded the orientation of wild-caught monarchs in southwestern Ontario and found that individuals generally flew in a southwest direction toward the wintering grounds. When displaced 2,500 km to the west, the same individuals continued to fly in a general southwest direction, suggesting that monarchs use a simple vector-navigation strategy (i.e., use a specific compass bearing without compensating for displacement). Using over 5 decades of field data, we also show that the directional concentration and the angular SD of recoveries from tagged monarchs largely conformed to two mathematical models describing the directional distribution of migrants expected under a vector-navigation strategy. A third analysis of tagged recoveries shows that the increasing directionality of migration from north to south is largely because of the presence of geographic barriers that guide individuals toward overwintering sites. Our work suggests that monarchs breeding in eastern North America likely combine simple orientation mechanisms with geographic features that funnel them toward Mexican overwintering sites, a remarkable achievement considering that these butterflies weigh less than a gram and travel thousands of kilometers to a site they have never seen.

To test if migratory monarch butterflies use polarized light patterns as part of their time-compensated sun compass, we recorded their virtual flight paths in a flight simulator while the butterflies were exposed to patches of naturally polarized blue sky, artificial polarizers or a sunny sky. In addition, we tested butterflies with and without the polarized light detectors of their compound eye being occluded. The monarchs' orientation responses suggested that the butterflies did not use the polarized light patterns as a compass cue, nor did they exhibit a specific alignment response towards the axis of polarized light. When given direct view of the sun, migratory monarchs with their polarized light detectors painted out were still able to use their time-compensated compass: non-clockshifted butterflies, with their dorsal rim area occluded, oriented in their typical south-southwesterly migratory direction. Furthermore, they shifted their flight course clockwise by the predicted approximately 90 degrees after being advance clockshifted 6 h. We conclude that in migratory monarch butterflies, polarized light cues are not necessary for a time-compensated celestial compass to work and that the azimuthal position of the sun disc and/or the associated light-intensity and spectral gradients seem to be the migrants' major compass cue.