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Journal of Experimental Biology partnership with Dryad

Water surface locomotion in tropical canopy ants
S. P. Yanoviak, D. N. Frederick


Upon falling onto the water surface, most terrestrial arthropods helplessly struggle and are quickly eaten by aquatic predators. Exceptions to this outcome mostly occur among riparian taxa that escape by walking or swimming at the water surface. Here we document sustained, directional, neustonic locomotion (i.e. surface swimming) in tropical arboreal ants. We dropped 35 species of ants into natural and artificial aquatic settings in Peru and Panama to assess their swimming ability. Ten species showed directed surface swimming at speeds >3 body lengths s−1, with some swimming at absolute speeds >10 cm s−1. Ten other species exhibited partial swimming ability characterized by relatively slow but directed movement. The remaining species showed no locomotory control at the surface. The phylogenetic distribution of swimming among ant genera indicates parallel evolution and a trend toward negative association with directed aerial descent behavior. Experiments with workers of Odontomachus bauri showed that they escape from the water by directing their swimming toward dark emergent objects (i.e. skototaxis). Analyses of high-speed video images indicate that Pachycondyla spp. and O. bauri use a modified alternating tripod gait when swimming; they generate thrust at the water surface via synchronized treading and rowing motions of the contralateral fore and mid legs, respectively, while the hind legs provide roll stability. These results expand the list of facultatively neustonic terrestrial taxa to include various species of tropical arboreal ants.


  • Author contributions

    S.P.Y. conceived the study, collected data, interpreted the results, conducted analyses, and wrote the manuscript. D.N.F. designed experiments, collected data, interpreted the results and wrote the manuscript.

  • Competing interests

    The authors declare no competing financial interests.

  • Funding

    This research was supported by the National Science Foundation [IOS-0843120 and DEB-1252614 to S.P.Y.].

  • Supplementary material

    Supplementary material available online at

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