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. 2018 Sep 4;9:3590. doi: 10.1038/s41467-018-05562-1

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© The Author(s) 2018

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Fig. 5 — Bat orientation behavior consists of two distinct modes—navigation and maneuvering—that are characterized by different angular speeds. a Left, a typical night-flight of an Egyptian fruit bat from its roosting cave (green ellipse) to distal foraging sites; scale bar, 2 km. Right, a zoom-in on the natural orientation behavior, showing epochs of navigation (commuting)—characterized by relatively straight flights during which there was little modulation of heading-direction, interspersed by periods of intensive maneuvering around fruit-trees (indicated by white arrows)—characterized by rapid turns (imagery produced using desktop version of Google Earth Pro). b Distribution of the combined (azimuth × pitch) angular velocity versus horizontal displacement of the bat. Maneuvering mode and navigation mode were classified according to the threshold (vertical dashed black line) at the minimum of the marginal distribution of the horizontal displacement. The marginal distribution of combined angular velocity is shown for all data (grey), and separately for navigation (red) and maneuvering modes (blue). c Angular-velocity distribution computed separately for pitch versus azimuth during navigation (left) and maneuvering (right). Inset (middle) shows the angular velocity distribution for the entire session. Note that during maneuvering (right), angular velocities in azimuth and pitch were correlated, suggesting that bats’ maneuvers were composed of rapid rotations in both azimuth and pitch. The data in b and c were pooled over all bats and flights that we recorded (45 bats with one nightly flight for each bat²⁶)