Behavioral response to conspecific songs on foraging territories of the heart-nosed bat

Abstract

Singing to create and defend territory boundaries is common among birds but rare in mammals. The African heart-nosed bat, Cardioderma cor, is hypothesized to use loud, low-frequency songs to reestablish foraging territories nightly. Territoriality can be defined ecologically, whereby an individual exclusively uses an area repeatedly, and behaviorally, through defense of an area. C. cor males sing on tightly abutting, exclusive areas nightly, which they return to throughout the season and sometimes across seasons. C. cor meets the ecological determinants of territoriality, but assessments of the use of song to maintain territories are lacking. We explore the singing behavior in this species by conducting song playback experiments within the borders of singing areas of 10 target individuals, with echolocation playbacks conducted as a control. In addition, we further explore the influence of song metrics on the behavioral response. Song playbacks prompted investigative and aggressive behavior, including passing by, approaching, and in one case, attacking the speaker, whereas echolocation did not. Additional post hoc analyses suggested that major song parameters, including song length, syllable frequency, intersyllable interval, and the number of double syllables comprising song stimuli influenced the level of response. For five bats we assessed whether their songs changed in response to the playback, and found that they sang faster, lower-frequency songs. These results are consistent with observations in other territorial animals including birds and gibbons, and provide a basis for further exploration of the territory defense hypothesis in the heart-nosed bat. We conclude that C. cor song features likely play an important role in mediating behavioral interactions within signaling networks of foraging bats.

Significance statement

It has been hypothesized that birdsong first evolved in support of territorial defense because it offered a cost-effective alternative to patrolling large spaces by flight. Singing-like behaviors have also been documented in several species of bats but never as a tool for maintaining foraging territories. However, evidence of foraging territoriality is scarce for bats, likely due to technical challenges associated with documenting such behaviors for a small, flying animal that may travel large distances at night. Here, we show for the first time that a bat responds to conspecific songs in a manner strikingly similar to many songbirds, providing support from outside songbirds for the hypothesis that territorial defense is a key selective pressure for singing in small, flying animals. This work provides the important basis for continuing to explore the role of singing, including song variability, in natural bat behavior outside of the roost.

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Acknowledgements

We thank the reviewers for their critical assessments of this work. We feel that their suggestions have greatly improved this manuscript. We thank Felix Mpelembwa, Nuhu Bahaty Mhapa, Alfred Absolem Mollel, and Nickodemasy Obeid for their assistance in field work. We thank the officers of Kikavuchini, Mkalama, and Longoi Villages, the Machame Weru Weru Ward, and the Hai District for their cooperation. Dassa Nkini of the Tanzania Conservation Resource Centre assisted with permit acquisition. We thank Brian Pierce, Thomas DeWitt, David Jones, Mirjam Knörnschild, and Kirsten Bohn for advice and discussion on statistics and experimental design. This research was funded by the National Science Foundation Graduate Research Fellowship Program, Bat Conservation International, and Wildlife Acoustics.

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Correspondence to Grace C. Smarsh.

Ethics declarations

All applicable international, national, and institutional guidelines for the use of animals were followed. We acquired all necessary permits and permissions to work with this species and in these regions: Institutional Animal Care and Use Committee, AUP 2012-087; Tanzania Commission for Science and Technology, 2014-53-ER-2012-58, 2013-65-NA-2012-58, and NA-2012-58.

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The authors declare that they have no conflict of interest.

Data availability

The datasets analyzed from the current study are available from the corresponding author on reasonable request.

Additional information

Communicated by M. Knoernschild

Electronic supplementary material

Online Resource 1

Video of a typical Cardioderma cor male singing in a perch. Individuals rotate while perched, broadcasting songs in bouts orally throughout the night (WMV 711 kb)

Online Resource 2

Spectrogram of an example contact call of C. cor, produced by both males and females while foraging. Contact calls consist of varying numbers of “upsweep” syllables (GIF 34 kb)

High resolution image (EPS 995 kb)

Online Resource 3

Song playlist for playback experiments. Each playlist consisted of a song repeated 20 times to create a set, which was repeated twice (Set 1 and Set 2) with 1 min of silence in between. Precontrol (PreC) and Postcontrol (PostC) time periods consisted of 5 min of silence at the beginning and the end of the playlist (GIF 8 kb)

High resolution image (EPS 686 kb)

Online Resource 4

Audio file of an example of a song playlist for playback. The silent PreC and PostC time periods on the ends of this playlist are five minutes in duration each (MP2 22126 kb)

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Smarsh, G.C., Smotherman, M. Behavioral response to conspecific songs on foraging territories of the heart-nosed bat. Behav Ecol Sociobiol 71, 142 (2017). https://doi.org/10.1007/s00265-017-2370-9

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Keywords

  • Bats
  • Singing behavior
  • Territoriality
  • Acoustic playback experiments
  • Cardioderma cor