Concatenation of 'alert' and 'identity' segments in dingoes' alarm calls.

Multicomponent signals can be formed by the uninterrupted concatenation of multiple call types. One such signal is found in dingoes, Canis familiaris dingo. This stereotyped, multicomponent 'bark-howl' vocalisation is formed by the concatenation of a noisy bark segment and a tonal howl segment. Both segments are structurally similar to bark and howl vocalisations produced independently in other contexts (e.g. intra- and inter-pack communication). Bark-howls are mainly uttered in response to human presence and were hypothesized to serve as alarm calls. We investigated the function of bark-howls and the respective roles of the bark and howl segments. We found that dingoes could discriminate between familiar and unfamiliar howl segments, after having only heard familiar howl vocalisations (i.e. different calls). We propose that howl segments could function as 'identity signals' and allow receivers to modulate their responses according to the caller's characteristics. The bark segment increased receivers' attention levels, providing support for earlier observational claims that barks have an 'alerting' function. Lastly, dingoes were more likely to display vigilance behaviours upon hearing bark-howl vocalisations, lending support to the alarm function hypothesis. Canid vocalisations, such as the dingo bark-howl, may provide a model system to investigate the selective pressures shaping complex communication systems.

The bark, the howl and the bark-howl: Identity cues in dingoes' multicomponent calls.

Dingoes (genus Canis) produce a stereotyped bark-howl vocalisation, which is a unimodal complex signal formed by the concatenation of two call types (a bark and a howl). Bark-howls may function as alarm signals, although there has been no empirical investigation of this vocalisation's structure or function. We quantified the content and efficacy of the bark and howl segments separately and when combined, using 140 calls from 10 individuals. We found that both segments are individually distinctive, although howl segments are more accurately classified, suggesting a higher level of individuality. Furthermore, howls convey signature characteristics that are conserved across different contexts of production, and thus may act as 'identity signals'. The individual distinctiveness of full bark-howls increases above that of isolated segments, which may be a result of selection on improved signal discriminability. Propagation tests revealed that bark-howls are best described as medium-range signals, with both segments potentially allowing for individual discrimination up to 200m regardless of environmental conditions. We discuss our findings regarding the fitness benefits of encoding identity cues in a potential alarm call and propose additional hypotheses for the function(s) of bark and howl segments.

Aggressive Bimodal Communication in Domestic Dogs, Canis familiaris.

Evidence of animal multimodal signalling is widespread and compelling. Dogs' aggressive vocalisations (growls and barks) have been extensively studied, but without any consideration of the simultaneously produced visual displays. In this study we aimed to categorize dogs' bimodal aggressive signals according to the redundant/non-redundant classification framework. We presented dogs with unimodal (audio or visual) or bimodal (audio-visual) stimuli and measured their gazing and motor behaviours. Responses did not qualitatively differ between the bimodal and two unimodal contexts, indicating that acoustic and visual signals provide redundant information. We could not further classify the signal as 'equivalent' or 'enhancing' as we found evidence for both subcategories. We discuss our findings in relation to the complex signal framework, and propose several hypotheses for this signal's function.

Mother knows best: functionally referential alarm calling in white-tailed ptarmigan.

Functionally referential alarm calls have stimulus specificity, distinct acoustic structure, and elicit different escape responses that are appropriate to the threat. The mechanisms by which escape responses are evoked are not fully understood and may range from eliciting innate responses to conveying representational information. White-tailed ptarmigan (Lagopus leucurus) are a long-lived alpine tundra grouse, which are preyed upon by aerial and terrestrial predators. We investigated the hypothesis that alarm calls of ptarmigan hens with chicks are functionally referential. We recorded hens' alarm calls in response to naturally occurring and model predators in California's Sierra Nevada alpine tundra for two summer seasons. We conducted playback experiments in the field to determine chick responses to alarm calls. Alarm calls commenced with an extended 'alerting' note followed by a series of staccato notes grouped into elements. Fundamental and dominant frequencies in element notes were significantly higher in terrestrial compared to aerial threat alarm calls. Playbacks of terrestrial threat alarm calls elicited an upright/alert position by chicks (75 % of responses). In response to aerial threat alarm call playbacks, chicks flattened to the ground and froze (80 % of responses). To our knowledge, this study provides the first empirical evidence of functionally referential alarm calling, including the responses of the receivers, in an avian species in the wild.