Free-ranging bats combine three different cognitive processes for roost localization.

Animals have evolved different cognitive processes to localize crucial resources that are difficult to find. Relevant cognitive processes such as associative learning and spatial memory have commonly been studied in a foraging related context under controlled laboratory conditions. However, in natural environments, animals can use multiple cognitive processes to localize resources. In this field study, we used a pairwise choice experiment and automatic roost monitoring to assess how individually marked, free-ranging Bechstein's bats belonging to two different colonies use associative learning, spatial memory and social information when localizing suitable day roosts. To our knowledge, this study tests for the first time how associative learning, spatial memory and social information are used in the process of roost localization in bats under the natural conditions. We show that, when searching for new roosts, bats used associative learning to discriminate between suitable and unsuitable roosts. For re-localizing previously occupied roosts, bats used spatial memory rather than associative learning. Moreover, bats significantly improved the localization of suitable unfamiliar roosts and tended to increase their accuracy to re-localize previously occupied day roosts using social information. Our field experiments suggest that Bechstein's bats make hierarchical use of different cognitive processes when localizing day roosts. More generally, our study underlines that evaluating different cues under natural conditions is fundamental to understanding how natural selection has shaped the cognitive processes used for localizing resources.