SINGING FOR YOUR SUPPER: SENTINEL CALLING BY KLEPTOPARASITES CAN MITIGATE THE COST TO VICTIMS

Despite the prevalence of vocal mimicry in animals, few functions for this behaviour have been shown. I propose a novel hypothesis that false mimicked alarm calls could be used deceptively to scare other species and steal their food. Studies have previously suggested that animals use their own species-specific alarm calls to steal food. However none have shown conclusively that these false alarms are deceptive, or that mimicked alarm calls are used in this manner. Here, I show that wild fork-tailed drongos (Dicrurus adsimilis) make both drongo-specific and mimicked false alarm calls when watching target species handling food, in response to which targets flee to cover abandoning their food. The drongo-specific and mimicked calls made in false alarms were structurally indistinguishable from calls made during true alarms at predators by drongos and other species. Furthermore, I demonstrate by playback experiments that two of these species, meerkats (Suricata suricatta) and pied babblers (Turdoides bicolor), are deceived by both drongo-specific and mimicked false alarm calls. These results provide the first conclusive evidence that false alarm calls are deceptive and demonstrate a novel function for vocal mimicry. This work also provides valuable insight into the benefits of deploying variable mimetic signals in deceptive communication.

Investigating a link between bill morphology, foraging ecology and kleptoparasitic behaviour in the fork-tailed drongo

Kleptoparasitism is a tactic used to acquire food opportunistically and has been shown to provide several benefits, including greater food intake rate and the acquisition of items not normally available during self-foraging. Host individuals may differ in their ability to defend themselves against kleptoparasitic attacks and therefore identifying those host individuals that are particularly vulnerable to attack could both provide energetic benefits and increase the efficiency of kleptoparasitism as a foraging strategy. Here, we show that the kleptoparasitic fork-tailed drongo (Dicrurus adsimilis) specifically targets juveniles when following groups of cooperatively breeding pied babblers (Turdoides bicolor). Drongos give alarm calls upon sighting a predator, thus providing extra predator vigilance to foraging pied babblers. However, drongos also use alarm calls to steal food items. During kleptoparasitic attacks, drongos give false alarm calls and then swoop down to steal food items dropped by alarmed babblers. Juvenile pied babblers are particularly vulnerable to attack because they (a) spend a longer period handling prey items prior to consumption and (b) respond to alarm calls primarily by immediately moving to cover, in contrast to adults who respond by looking up and visually scanning the surrounding area. Drongos attack juvenile babblers significantly more often than adults, with attacks on juveniles more likely to result in the successful procurement of a food item. This patterns of attack suggests that drongos are able to differentiate between individuals of different age when targeting pied babblers, thus increasing the efficiency of kleptoparasitism as a foraging strategy.

In many cases of interspecific kleptoparasitism, hosts develop defensive behaviors to minimize the impact of kleptoparasites. Because vigilance and defensive behaviors are often costly, selection should favor hosts that adjust the amount of investment needed to minimize losses to kleptoparasitism. However, examples of such facultative responses are rare. Here, we investigate the response of cooperatively breeding pied babblers (Turdoides bicolor) to the drongo (Dicrurus adsimilis), an avian kleptoparasite that regularly follows pied babbler groups, often giving alarm calls to alert the group to predators but also occasionally giving false alarm calls in order to steal food items. We show that pied babbler response to drongos varies markedly according to babbler group size. In small groups, where there are few individuals available to act as sentinels, babblers sentinel less when a drongo is present and respond strongly to drongo alarm calls. However, in large groups, where there are many individuals available to participate in predator vigilance, babblers sentinel more often when a drongo is present, rarely respond to drongo alarm calls, and aggressively displace drongos, with a consequent decline in the number of successful kleptoparasitism events. This behavior represent a facultative response to a kleptoparasite according to the costs versus benefits of tolerating their presence.

Audience effects are increasingly recognized as an important aspect of intraspecific communication. Yet despite the common occurrence of interspecific interactions and considerable evidence that individuals respond to the calls of heterospecifics, empirical evidence for interspecific audience effects on signalling behaviour is lacking. Here we present evidence of an interspecific audience effect on the alarm-calling behaviour of the kleptoparasitic fork-tailed drongo (Dicrurus adsimilis). When foraging solitarily, drongos regularly alarm at aerial predators, but rarely alarm at terrestrial predators. In contrast, when drongos are following terrestrially foraging pied babblers (Turdoides bicolor) for kleptoparasitic opportunities, they consistently give alarm calls to both aerial and terrestrial predators. This change occurs despite no difference in the amount of time that drongos spend foraging terrestrially. Babblers respond to drongo alarm calls by fleeing to cover, providing drongos with opportunities to steal babbler food items by occasionally giving false alarm calls. This provides an example of an interspecific audience effect on alarm-calling behaviour that may be explained by the benefits received from audience response.

Vocal recognition of former group members, but not unknown kin, in the cooperatively breeding southern pied babbler

In risky environments, where threats are unpredictable and the quality of information about threats is variable, all individuals face two fundamental challenges: balancing vigilance against other activities, and determining when to respond to warning signals. The solution to both is to obtain continuous estimates of background risk, enabling vigilance to be concentrated during the riskiest periods and informing about the likely cost of ignoring warnings. Human surveillance organizations routinely produce such estimates, frequently derived from indirect cues. Here we show that vigilant individuals in an animal society (the pied babbler, Turdoides bicolor) perform a similar role. We ask (i) whether, in the absence of direct predator threats, pied babbler sentinels react to indirect information associated with increased risk and whether they communicate this information to group mates; (ii) whether group mates use this information to adjust their own vigilance, and whether this influences foraging success; and (iii) whether information provided by sentinels reduces the likelihood of inappropriate responses to alarm calls. Using playback experiments, we show that: (i) sentinels reacted to indirect predator cues (in the form of heterospecific alarm calls) by giving graded surveillance calls; (ii) foragers adjusted their vigilance in reaction to changes in surveillance calls, with substantial effects on foraging success; and (iii) foragers reduced their probability of responding to alarm calls when surveillance calls indicated lowered risk. These results demonstrate that identifying attacks as they occur is only part of vigilance: equally important is continuous surveillance providing information necessary for individuals to make decisions about their own vigilance and evasive action. Moreover, they suggest that a major benefit of group living is not only the increased likelihood of detecting threats, but a marked improvement in the quality of information available to each individual.

Recruitment Calling: A Novel Form of Extended Parental Care in an Altricial Species

Kleptoparasitism by attacks versus false alarm calls in fork-tailed drongos

Acoustic communication plays a vital role in passing or sharing information between individuals. Identifying the biological meaning of vocal signals is crucial in understanding the survival strategies of animals. However, there are many challenges in identifying the true meaning of such signals. The plateau pika (Ochotona curzoniae) is a call-producing mammal endemic to the Qinghai-Tibet plateau (QTP) and considered a keystone species owing to its multiple benefits in alpine rangeland ecosystems. Previous studies have shown that plateau pikas emit alarm calls as part of their daily activities. However, only field observations have been used to identify these alarm calls of the plateau pika, with no attempts at using playback experiments. Here, we report the alarm calling of plateau pikas through field observations as well as simulated predator and playback experiments in the Eastern QTP from 2021 to 2022. We found that both female and male adults emitted alarm calls, the signals of which comprised only one syllable, with a duration of 0.1-0.3 s. There were no differences in the characteristics between the observed alarm calls and those made in response to the simulated predator. The duration of the alarm call response varied with altitude, with plateau pikas living at higher altitudes responding at shorter durations than those at lower altitudes.

Social learning is beneficial in almost every domain of a social animal's life, but it is particularly important in the context of predation and foraging. In both contexts, social animals tend to produce acoustically distinct vocalizations, alarms, and food calls, which have remained somewhat of an evolutionary conundrum as they appear to be costly for the signaller. Here, we investigated the hypothesis that food calls function to direct others toward novel food items, using a playback experiment on a group of chimpanzees. We showed chimpanzees novel (plausibly edible) items while simultaneously playing either conspecific food calls or acoustically similar greeting calls as a control. We found that individuals responded by staying longer near items previously associated with food calls even in the absence of these vocalizations, and peered more at these items compared with the control items, provided no conspecifics were nearby. We also found that once chimpanzees had access to both item types, they interacted more with the one previously associated with food calls than the control items. However, we found no evidence of social learning per se. Given these effects, we propose that food calls may gate and thus facilitate social learning by directing listeners' attention to new feeding opportunities, which if integrated with additional cues could ultimately lead to new food preferences within social groups.

Animals resolve conflicts over the share of resources by competing physically or signalling motivation with honest signals of need. In some species, young siblings vocally signal to each other their hunger level and the most vocal individual deters its siblings from competing for the non-divisible food item delivered at the next parental visit. This so-called sibling negotiation for forthcoming food has been studied only in this context. It therefore remains unclear whether siblings could also negotiate access to a pool of divisible resources, a situation that is similar to a group of individuals competing for an accessible food resource. To tackle this issue, we placed barn owl (Tyto alba) nestlings singly in artificial nests containing several mice, and we simulated the presence of a sibling calling at low or high rate using playback experiments. If nestling barn owls vocally negotiate over a divisible food stock, we propose the following two predictions. First, nestlings would vocally signal before eating from this stock of food, and second, numerous playback vocalizations would inhibit feeding. Accordingly, singleton nestlings vocalized just before consuming food stored in their artificial nest and they delayed the consumption of the food stock if hearing many playback calls. The production of such food-associated vocalizations has been observed in foraging adults in various birds and mammals, but never in young animals and when resource is divisible and easily accessible. Our study raises the possibility that vocal communication could evolve in a variety of competitive contexts. We present here the first experimental evidence that sibling barn owls use food-associated vocalizations to compete over the preys stored in the nest. Owlets emit calls just before consuming an available food item and broadcasting calls induces nestlings to temporarily refrain from eating from the food stock. This raises the possibility that vocal communication can mediate the share of a food stock accessible to all competitors.

Possible use of heterospecific food-associated calls of macaques by sika deer for foraging efficiency

We report on a simple and practical application of HARK, an easily available and portable system for bird song localization using an open-source software for robot audition HARK, to a deeper understanding of ecoacoustic dynamics of bird songs, focusing on a fine-scaled temporal analysis of song movement — song type dynamics in playback experiments. We extended HARKBird and constructed a system that enables us to conduct automatic playback and interactive experiments with different conditions, with a real-time recording and localization of sound sources. We investigate how playback of conspecific songs and playback patterns can affect vocalization of two types of songs and spatial movement of an individual of Japanese bush-warbler, showing quantitatively that there exist strong relationships between song type and spatial movement. We also simulated the ecoacoustic dynamics of the singing behavior of the focal individual using a software, termed Bird song explorer, which provides users a virtual experience of acoustic dynamics of bird songs using a 3D game platform Unity. Based on experimental results, we discuss how our approach can contribute to ecoacoustics in terms of two different roles of sounds: sounds as tools and subjects.

Close calling regulates spacing between foraging competitors in the group-living pied babbler