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Recognition of familiarity on the basis of howls: a playback experiment in a captive group of wolves
in BehaviourPlayback experiments were conducted with a pack of captive Iberian wolves. We used a habituation–discrimination paradigm to test wolves’ ability to discriminate howls based on: (1) artificial manipulation of acoustic parameters of howls and (2) the identity of howling individuals. Manipulations in fundamental frequency and frequency modulation within the natural range of intra-individual howl variation did not elicit dishabituation, while manipulation of modulation pattern did produce dishabituation. With respect to identity, across trials wolves habituated to unfamiliar howls by a familiar wolf (i.e., no direct contact, but previous exposure to howls by this wolf), but not to unfamiliar howls from unfamiliar wolves (i.e., no direct contact and no previous exposure to howls by these wolves). Modulation pattern seems to be an important bioacoustic feature for individual recognition. Overall, our results provide the first experimental evidence that wolves can discriminate individuals based on the acoustic structure of their howls.
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Figures
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Acoustic stimuli created (upper panel) and playback protocol conducted (lower panel). ∗From our sample of recorded howls (emitted by six wolves unrelated to Senda Viva’s wolves) we randomly selected a wolf (Familiar Wolf) to create the habituation stimulus (Familiar Stimulus) to habituate the group of captive wolves to its howls. This figure is published in colour in the online edition of this journal, which can be accessed via
http://booksandjournals.brillonline.com/content/journals/1568539x . -
Familiar stimulus and playback stimuli created modifying the acoustic structure of howls. FS, Familiar Stimulus; FS+100, +200, −100 and −200, familiar stimulus increasing and decreasing the fundamental frequency by 100 or 200 Hz; FS+cofm, FS increasing the coefficient of frequency modulation; FS rev, FS reversing the howls.
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Wolf enclosure sketch. Z1 and Z2, zones where the wolves were visible from the observation point; Z3, area where wolves could not be observed; OP, observation point; S, speaker.
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Level of response (number of Attention displayed by all the wolves within sight) during the two min following the beginning of the stimulus. (Left) Level of response obtained to the Familiar Stimulus (FS) in the valid trials () during the habituation phase. (Right) Average level of response (mean ± SE number of Attention) for each stimulus taking into account all the valid trials. FS (d), FS during the habituation–dishabituation phase; FS+100, +200, −100, −200: FS adding or subtracting 100 or 200 Hz; FS+cofm, FS manipulating only the coefficient of frequency modulation; FS Rev, FS with the howls reversed; UFhFw, unfamiliar howls by the familiar wolf; UFhUFw, unfamiliar howls by unfamiliar wolves.
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Level of response (i.e., number of Attention; see Methods) of wolves to unfamiliar howls emitted by a familiar wolf or by unfamiliar wolves. Solid lines reflect predicted values for each treatment, while dashed lines show the standard error interval (GLM model: number of attention − treatment ∗ playback trial + group size; Poisson error distribution). Note that random jitter has been introduced in the x axis to distinguish data points from both treatments. This figure is published in colour in the online edition of this journal, which can be accessed via
http://booksandjournals.brillonline.com/content/journals/1568539x .
Acoustic stimuli created (upper panel) and playback protocol conducted (lower panel). ∗From our sample of recorded howls (emitted by six wolves unrelated to Senda Viva’s wolves) we randomly selected a wolf (Familiar Wolf) to create the habituation stimulus (Familiar Stimulus) to habituate the group of captive wolves to its howls. This figure is published in colour in the online edition of this journal, which can be accessed via
Familiar stimulus and playback stimuli created modifying the acoustic structure of howls. FS, Familiar Stimulus; FS+100, +200, −100 and −200, familiar stimulus increasing and decreasing the fundamental frequency by 100 or 200 Hz; FS+cofm, FS increasing the coefficient of frequency modulation; FS rev, FS reversing the howls.
Wolf enclosure sketch. Z1 and Z2, zones where the wolves were visible from the observation point; Z3, area where wolves could not be observed; OP, observation point; S, speaker.
Level of response (number of Attention displayed by all the wolves within sight) during the two min following the beginning of the stimulus. (Left) Level of response obtained to the Familiar Stimulus (FS) in the valid trials () during the habituation phase. (Right) Average level of response (mean ± SE number of Attention) for each stimulus taking into account all the valid trials. FS (d), FS during the habituation–dishabituation phase; FS+100, +200, −100, −200: FS adding or subtracting 100 or 200 Hz; FS+cofm, FS manipulating only the coefficient of frequency modulation; FS Rev, FS with the howls reversed; UFhFw, unfamiliar howls by the familiar wolf; UFhUFw, unfamiliar howls by unfamiliar wolves.
Level of response (i.e., number of Attention; see Methods) of wolves to unfamiliar howls emitted by a familiar wolf or by unfamiliar wolves. Solid lines reflect predicted values for each treatment, while dashed lines show the standard error interval (GLM model: number of attention − treatment ∗ playback trial + group size; Poisson error distribution). Note that random jitter has been introduced in the x axis to distinguish data points from both treatments. This figure is published in colour in the online edition of this journal, which can be accessed via
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