Social rules govern vocal competition in the barn owl

To resolve the share of limited resources, animals often compete through exchange of signals about their relative motivation to compete. When two competitors are similarly motivated, the resolution of conflicts may be achieved in the course of an interactive process. In barn owls, Tyto alba, in which siblings vocally compete during the prolonged absence of parents over access to the next delivered food item, we investigated what governs the decision to leave or enter a contest, and at which level. Siblings alternated periods during which one of the two individuals vocalized more than the other. Individuals followed turn-taking rules to interrupt each other and momentarily dominate the vocal competition. These social rules were weakly sensitive to hunger level and age hierarchy. Hence, the investment in a conflict is determined not only by need and resource-holding potential, but also by social interactions. The use of turn-taking rules governing individual vocal investment has rarely been shown in a competitive context. We hypothesized that these rules would allow individuals to remain alert to one another's motivation while maintaining the cost of vocalizing at the lowest level

Information retention during competitive interactions : siblings need to constantly repeat vocal displays

Memory is essential to adjust behaviour according to past experience. In societies where animals interact on numerous occasions, memory of previous social interactions may help optimise investment in competition. How long information about the resource holding potential and motivation to compete of conspecifics is retained depends on how fast the value of this information fades, but also on the cost and benefit of retaining information. Information retention has never been investigated in the context of interactions prevailing within the family and more specifically sibling competition. In the absence of parents, barn owl (Tyto alba) nestlings vocally compete for priority of access to the next indivisible food item brought by a parent. The finding that owlets eavesdrop on vocal interactions between siblings to adjust investment in vocalization once competing with them suggests that they memorize siblings' vocal interactions. Playback experiments showed that owlets take into account the past siblings' vocal performance that signals hunger for at least 15 min, but only if the performance was witnessed during a sufficiently long period of time (30 min). Moreover, using natural vocal exchanges in another set of individuals, we showed that sibling signalling was no more taken into account after a few minutes. This suggests that young barn owls need to continuously display their motivation to trigger siblings' withdrawal from the current competition. Repeating a vocal display may ensure its honesty. Studying the extent to which individuals retain past information is important to understand how individuals adjust their competitive investment over resources

Big brother is watching you: eavesdropping to resolve family conflicts

Adult animals can eavesdrop on behavioral interactions between potential opponents to assess their competitive ability and motivation to contest resources without interacting directly with them. Surprisingly, eavesdropping is not yet considered as an important factor used to resolve conflicts between family members. In this study, we show that nestling barn owls (Tyto alba) competing for food eavesdrop on nestmates' vocal interactions to assess the dominance status and food needs of opponents. During a first training playback session, we broadcasted to singleton bystander nestlings a simulated vocal interaction between 2 prerecorded individuals, 1 relatively old (i.e., senior) and 1 younger nestling (i.e., junior). One playback individual, the "responder," called systematically just after the "initiator" playback individual, hence displaying a higher hunger level. To test whether nestlings have eavesdropped on this interaction, we broadcasted the same prerecorded individuals separately in a subsequent playback test session. Nestlings vocalized more rapidly after former initiators' than responders' calls and they produced more calls when the broadcasted individual was formerly a junior initiator. They chiefly challenged vocally juniors and initiators against whom the likelihood of winning a vocal contest is higher. Owlets, therefore, identified the age hierarchy between 2 competitors based on their vocalizations. They also memorized the dynamics of competitors' previous vocal interactions, and used this information to optimally adjust signaling level once interacting with only 1 of the competitor. We conclude that siblings eavesdrop on one another to resolve conflicts over parental resources

Nestling barn owls assess short-term variation in the amount of vocally competing siblings.

Assessing the amount of rivals is crucial to optimally adjust investment into a contest. If laboratory animals show numerical abilities, little is known about the ecological implications particularly in young animals. The two to nine barn owl (Tyto alba) siblings vocally compete for priority of access to food resources before parents actually deliver them. In dyads, the individual that vocalizes at the highest rate in the absence of parents deters its siblings from competing for next delivered prey. We tested the novel hypothesis that to optimally adjust vocal investment, barn owl nestlings assess how many of their siblings are currently competing. To singleton owlets, we broadcasted a fixed global number of calls emitted by one, two or four pre-recorded unfamiliar nestlings. We could thus distinguish the independent effect on singletons' vocal behavior of the global number of calls produced by a brood from the number of competitors that produced these calls. Overall, nestlings retreated more from vocal contest when facing more competitors. However, in front of one highly motivated competitor, nestlings refrained from vocalizing to a larger extent than when competing against more but less motivated individuals. Therefore, young animals assess variation in the number of currently competing siblings based on individual-specific vocal cues

Dominant nestlings displaying female-like melanin coloration behave altruistically in the barn owl

When competing over parental resources, young animals may be typically selfish to the point of siblicide. This suggests that limited parental resources promote the evolution of sibling competition rather than altruistic or cooperative behaviours. In striking contrast, we show here that in 71% of experimental three-chick broods, nestling barn owls, Tyto alba, gave food to their siblings on average twice per night. This behaviour prevailed in the first-born dominant nestlings rather than the last-born subordinate nestlings. It was also more prevalent in individuals displaying a heritable dark phaeomelanin-based coloration, a typical female-specific plumage trait (owls vary from dark reddish to white, females being on average darker reddish than males). Stealing food items from siblings, which occurred in 81% of the nests, was more frequent in light than dark phaeomelanic dominant nestlings. We suggest that food sharing has evolved in the barn owl because parents store prey items in their nest that can be used by the offspring to feed their nestmates to derive indirect (kin selection) or direct benefits (pseudoreciprocity or by-product mutualism). The cost of feeding siblings may be relatively low for dominant individuals while the indirect genetic benefits could be high given that extrapair paternity is infrequent in this species. Thus, in situations in which young animals have access to more food resources than they currently need, they can altruistically share them with their siblings

Responsiveness to siblings' need increases with age in vocally negotiating barn owl nestlings

In animal societies, individuals should optimize the way they behave in relation to the behavior displayed by their conspecifics. This social competence, i.e., the ability to adjust behavior to the social context, can vary between individuals, but also improve with age and experience. This aspect, although important, has rarely been studied. We tested whether the ability to adjust behavior to siblings develops with age in barn owl nestlings (Tyto alba). In this species, young siblings show intense social interactions referred to as Bsibling negotiation. Indeed, because parents bring a single indivisible food item at each visit to the nest, all the effort invested in sibling competition is only paid back in the nestling that is able to monopolize the food item. Therefore, before the arrival of parents, siblings vocally inform each other about their relative hunger level so that they can optimally invest in sibling competition, with the most vocal, and hence hungry, nestling momentarily deterring its siblings from competing. This process implies that siblings have the ability to adjust their behavior in relation to the behavior of their siblings, a process that could change with age. In a series of experiments, we examined how nestlings of different ages respond to the vocal behavior of siblings. We show here that older nestlings adjusted their vocal behavior more finely than younger nestlings in relation to the behavior of their siblings. Elders also more readily refrained from eating in front of a hungry sibling. These patterns could arise because owlets' social competence develops with age or because they adopt different competitive and cooperative strategies according to their age. Significance statement In sibling barn owls, competition for food brought by parents is settled by vocalization. Highly vocal owlets induce their siblings to call less and to let them eat in priority once parents are back with a prey item, a process referred to as Bsibling negotiation." Nestling barn owls adjust their investment in sibling competition according to two parameters: their hunger level and the vocal behavior of their siblings. We analyzed the relative importance of these two parameters in differently aged owlets. Younger owlets adjusted the intensity of vocalizing primarily in relation to their own hunger level, which was efficient in modifying older nestlings' behavior, as older nestlings readily withdrew from vocal contest and refrained from eating in front of highly vocal siblings. Hence, social adjustment changed with age in owlets, older ones being more sensitive to the signals of need of their siblings

No need to shout: Effect of signal loudness on sibling communication in barn owls Tyto alba

In animal communication, signal loudness is often ignored and seldom measured. We used a playback experiment to examine the role of vocal loudness (i.e., sound pressure level) in sibling to sibling communication of nestling barn owls Tyto alba. In this species, siblings vocally negotiate among each other for priority access to parental food resources. Call rate and call duration play key roles in this vocal communication system, with the most vocal nestlings deterring their siblings from competing for access to the food item next delivered by parents. Here, we broadcast calls at different loudness levels and call rate to live nestlings. The loudness of playback calls did not affect owlets' investment in call rate, call duration or call loudness. The rate at which playback calls were broadcast affected owlets' call rate but did not influence their response in terms of loudness. This suggests that selection for producing loud signals may be weak in this species, as loud calls may attract predators. Moreover, given that owlets do not overlap their calls and that they communicate to nearby siblings in the silence of the night, loud signals may not be necessary to convey reliable information about food need