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

Non-Vocal Behaviors Are More Frequent During the Decisive Negotiation Phases in Barn Owl Siblings.

Animals produce vibrations or noises by means of body movements, which can play a role in communication. These behaviors enhance signal transmission or receiver attention and could be specifically used during turn-taking phases of a reciprocal exchange of signals. In the barn owl Tyto alba, nestlings vocalize one after the other to negotiate which individual will have priority access to the impending prey item to be delivered by the parents. Owlets adjust their vocalization to their own hunger level and to their siblings' vocalization, withdrawing from the contest in front of highly vocal, and hence hungry, motivated nestmates. As sibling negotiation is a multicomponent display, we examined whether body movements could also be part of the negotiation process. To this end, we analyzed whether the vocalizations of one nestling affected its nestmate's movements in three separate experiments: in natural nests, in the lab, and using a playback procedure. Nestling barn owls move in a variety of ways, such as repeated tapping of the floor with a foot, scratching the floor with claws, or flapping wings. Body movements were more frequent during the turn-taking phases of vocal interactions, when siblings emitted longer calls and at a greater rate. Once an individual monopolized vocal activity, siblings became less vocal and less active. Moreover, owlets produced more noisy body movements during the phases of vocal interactions which are crucial to prevail in negotiation. Non-vocal physical activities might reinforce vocal signals during sibling to sibling (sib-sib) interactions, or reflect owlets' arousal, in the critical period during which they vocally settle which individual will dominate the competition

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

Sibling rivalry and vocal negotiation in the barn owl Tyto alba

Chez les animaux, les jeunes dépendant des parents durant leur développement sont en compétition pour obtenir la nourriture, qu'ils quémandent par des cris et postures ostentatoires et se disputent physiquement. Les frères et soeurs n'ont pas la même compétitivité, en particulier s'ils diffèrent en âge, et leur niveau de faim fluctue dans le temps. Comme dans tout type de compétition, chacun doit ajuster son investissement aux rivaux, c'est à dire aux besoins et comportements de ses frères et soeurs. Dans le contexte de la famille, selon la théorie de sélection de parentèle, les jeunes bénéficient de leur survie mutuelle et donc de la propagation de la part de gènes qu'ils ont en commun. L'hypothèse de la « négociation frères-soeurs » prédit que, sous certaines conditions, les jeunes négocient entre eux la nourriture, ce qui réduit les coûts de compétition et permet de favoriser les frères et soeurs les plus affamés. La littérature actuelle se focalise sur les signaux de quémande entre enfants et parents et les interactions compétitives frères-soeurs sont étudiées principalement au sein de paires, alors que les nichées ou portées en comprennent souvent de nombreux. Cette thèse vise à mieux comprendre comment et jusqu'à quel point plusieurs jeunes ajustent mutuellement leurs signaux de besoin. C'est une question importante, étant donné que cela influence la répartition de nourriture entre eux, donc la résolution du conflit qui les oppose et à terme leur valeur évolutive. Le modèle d'étude est la chouette effraie (Tyto alba), chez laquelle jusqu'à neufs poussins émettent des milliers de cris chacun par nuit. Ils négocieraient entre eux la prochaine proie indivisible rapportée au nid avant que les parents ne reviennent : un poussin affamé crie plus qu'un autre moins affamé, ce qui dissuade ce dernier de crier en retour et par la suite de quémander la nourriture aux parents. L'investissement optimal correspondrait donc à écarter son frère en permanence vu que l'arrivée des parents est imprévisible, mais à moindre coût. Dans un premier axe, nous avons exploré au sein de dyades les mécanismes acoustiques permettant aux poussins de doser leur effort vocal durant les heures de compétition où ils sont laissés seuls au nid. Nous avons trouvé que les poussins évitent de crier simultanément, ce qui optimiserait la discrimination du nombre et de la durée de leurs cris, lesquels reflètent de façon honnête leur niveau de faim et donc leur motivation. L'alternance des cris paraît particulièrement adaptée au fait que les poussins se fient à des variations temporelles subtiles dans le rythme et la durée de leurs vocalisations pour prendre la parole. En particulier, allonger ses cris tout en criant moins dissuade efficacement le rival de répondre, ce qui permet de monopoliser la parole dans de longs « monologues ». Ces règles seraient universelles puisqu'elles ne dépendent pas de la séniorité, de la faim, ni de la parenté et les poussins répondent à un playback de façon similaire à un vrai frère. Tous ces résultats apportent la première preuve expérimentale que les juvéniles communiquent de façon honnête sur leurs besoins, ajustent activement le rythme de leurs cris et utilisent des composantes multiples de leurs vocalisations d'une façon qui réduit le coût de la compétition. De plus, il s'agit de la première démonstration que des règles de conversation régissent de longs échanges vocaux chez les animaux de façon comparable aux règles basiques observées chez l'Homme. Dans un second axe, nous avons exploré les stratégies comportementales que les poussins adoptent pour rivaliser avec plusieurs frères et soeurs, par le biais d'expériences de playback. Nous avons trouvé que les poussins mémorisent des asymétries de compétitivité entre deux individus qui dialoguent et répondent plus agressivement au moins compétitif une fois qu'ils sont confrontés à chacun isolément. Dans la même ligne, quand ils entendent un nombre variable d'individus criant à un taux variable, les poussins investissent le plus contre des rivaux moins nombreux et moins motivés. En accord avec les prédictions des modèles théoriques, les poussins de chouette effraie escaladent donc les conflits pour lesquels leur chance de gagner contrebalance le plus l'énergie dépensée. Nous révélons ainsi que 1) les jeunes frères et soeurs 'espionnent' les interactions de leurs rivaux pour évaluer leur compétitivité relative, ce qui est sans doute moins coûteux qu'une confrontation directe avec chacun, et 2) dosent leur investissement vocal en fonction du nombre de rivaux actuellement en compétition et de leur motivation de façon concomitante. Ces résultats montrent que les interactions entre frères et soeurs au nid reposent sur des mécanismes similaires à ceux observés, mais encore de façon anecdotique, chez les adultes non apparentés qui se disputent les territoires et partenaires sexuels. Cette thèse souligne donc combien il est crucial de considérer dorénavant la famille comme un réseau de communication à part entière pour mieux comprendre comment les jeunes résolvent les conflits autour du partage des ressources parentales. Plus généralement, elle révèle l'importance de la dynamique temporelle des vocalisations dans les conflits et la communication des animaux. A la lumière de nos résultats, la chouette effraie apparaît comme un modèle clé pour de futures recherches sur la résolution des conflits et la communication acoustique. - In species with parental care, offspring contest priority access to food by begging through conspicuous postures and vocalisations and by physically jockeying. Siblings differ in their competitiveness, especially in the case of age and size hierarchies, and their hunger level fluctuates in time. As in competition in general, each individual should adjust its investment to opponents that is to say to its siblings' needs and behaviours. In the particular context of family, according to kin selection theory, siblings derive extra fitness benefits from their mutual survival and hence the spreading of the genes they share. The "sibling negotiation" predicts that, under certain conditions, young would negotiate among them priority access to food, which reduces competition costs and enables promoting the most hungry siblings. To date, the literature focuses on signals of need between parents and offspring and competitive interactions (in particular among siblings) are mostly studied within pairwise interactions, yet they commonly involve more numerous rivals. This PhD aims at better understanding how and the extent to which several young siblings compete through signalling. This is important since this influences how food is allocated among them, thus the outcome of sibling rivalry and ultimately their fitness. I use the barn owl (Tyto alba) as a model, in which the one to nine nestlings emit a simple noisy call thousands of times per night. Thereby, they would negotiate among them priority access to the indivisible food next delivered prior to parents' feeding visits. A hungry nestling emits more calls than a less hungry sibling, which deters it to call in return and ultimately beg food at parents. The optimal investment thus corresponds to constantly deterring the rival to compete, given that parents' arrival is unpredictable, but at the lowest costs. In the first axis of my thesis, we explored within dyads the acoustic mechanisms by which owlets dose vocal effort when competing during the hours they are left alone. We found that owlets avoid overlapping each other's calls. This would enhance the discrimination of both call number and duration, which honestly reflect individuals' hunger level and hence motivation to compete. Such antiphony seems best adapted to the fact that siblings actually use subtle temporal variations in the rhythm and duration of their calls to take or give their turn. Owlets alternate monologs, in which lengthening calls efficiently deters the rival to respond while reducing call number. Such rules depend neither on seniority, hunger level nor kinship since nestlings responded similarly to a live sibling and an unrelated playback individual. Taken together, these findings provide the first experimental proof that dependent young honestly communicate about their need, actively adjust the timing of their calls and use multicomponent signals in a way that reduces vocal costs. Moreover, this is the first demonstration of conversational rules underlying animal long-lasting vocal exchanges comparable to the basic turn-taking signals observed in humans. In the second axis, we focused on the behavioural strategies owlets adopt to compete with more than one sibling, using playback experiments. We found that singleton bystanders memorised competitive asymmetries between two playback individuals dialoguing and responded more aggressively to the submissive one once they later faced each of both alone. Moreover, when hearing a varying number of nestlings calling at varying rates, owlets vocally invested the most towards fewer and less motivated rivals. In line with predictions from models on conflict settlement, barn owls thus escalate contests in which their chance of winning best counterbalances the energy spent. These results reveal that young socially eavesdrop on their siblings' interactions to assess their relative competitiveness at likely lower costs than direct confrontation, and dose vocal effort relative to both their number and motivation. This shows that young siblings' interactions imply mechanisms similar to those observed, yet still anecdotally, in unrelated adults that contest mates and territories. This PhD therefore highlights how crucial it is to further consider family as a communication network to better understand how siblings resolve conflicts over the share of parental resources. More generally, it provides important insights into the role of the temporal dynamics of signalling during animal contests and communication. In the light of our findings, the barn owl emerges as a key model for future research on conflict resolution and acoustic communication in animals

Contribution aux méthodes de dosage du cobalt, du cuivre, du nickel et du vanadium dans les roches silicatées

Exposé d'unie méthode spectrographique de dosage du cobalt, du chrome, du cuivre, du nickel et du vanadium dans des roches silicatées de compositions variées, après attaque sulfofluorhydrique de l'échantillon, par la technique des additions successives. Étude de la reproductibilité des dosages, et comparaison avec les résultats obtenus par une méthode indépendante, la colorimétrie. Discussion des teneurs déterminées pour les échantillons standards américains, le granite G₁ et la diabase W₁.Maurel Pierre, Ruppli C. Contribution aux méthodes de dosage du cobalt, du cuivre, du nickel et du vanadium dans les roches silicatées. In: Bulletin de la Société française de Minéralogie et de Cristallographie, volume 84, 4, 1961. pp. 363-376

Réaction du granite fondu sous pression d'eau avec la calcite et une argile. Application au métamorphisme de contact

Étude de la réaction de petits cylindres de CaCO₃ et d'argile immergés dans un magma granitique fondu à 800° C, sous une pression de 1 800 bars. Dans le le cas du calcaire la réaction consiste en une migration massive de silice du magma vers CaCO₃ qui est converti en wollastonite. Dans le cas de l'argile, qui, dans ces conditions de température et de pression se liquéfie avec formation de cristaux de spinelle, ce liquide et le magma tendent vers une composition chimique commune par diffusion du sodium et du potassium et dissolution d'une partie du spinelle ; mais une composition chimique uniforme ne s'établit pas dans tout le système par suite de la persistance de cristaux de spinelle résiduels dans la région initialement occupée par l'argile. On observe en fin d'expérience, et éventuellement à l'équilibre, la persistance côte à côte de deux systèmes de compositions chimiques très différentes, séparés par une frontière nette.Wyart Jean, Ruppli C., Sabatier Germain. Réaction du granite fondu sous pression d'eau avec la calcite et une argile. Application au métamorphisme de contact. In: Bulletin de la Société française de Minéralogie et de Cristallographie, volume 83, 4-6, 1960. pp. 128-133

Sur une argile essentiellement formée de chlorite associée au gypse des environs d'Amélie-les-Bains

Romero Aristides, Ruppli C., Sabatier Germain. Sur une argile essentiellement formée de chlorite associée au gypse des environs d'Amélie-les-Bains. In: Bulletin de la Société française de Minéralogie et de Cristallographie, volume 84, 4, 1961. p. 404

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

Nocturnal, diurnal and bimodal patterns of locomotion, sibling interactions and sleep in nestling Barn Owls

Temporal variation in physical activity is mainly determined by the day-night cycle. While this may be true for diurnal species whose vision at night is often poor, the situation might be more complex in nocturnal animals as many such species can see both in the dark and in the daylight. We examined in Barn Owl (Tyto alba) nestlings whether temporal variation of behavioural activities and sleep is shaped by parental feeding visits occurring during the first part of the night and the extent to which they also occur during daylight hours. We measured several behaviours in 280 individuals from 90 broods recorded in 4 years. Parental feeding visits progressively declined in frequency from the beginning to the end of the night, and a number of offspring behaviours followed the same pattern of activity (feeding, vocalization and self-preening). Surprisingly, nestlings were awake not only at sunset, but also at sunrise. Several behaviours (locomotion, wing flapping and sibling interactions, such as pecking and allopreening among nestlings) showed peaks of activity at sunset and sunrise, suggesting that they were performed for other reasons than to interact with parents. Allopreening was performed more often during the day than at night. We conclude that although adult Barn Owls are nocturnal, nestlings display a complex temporal pattern of activity that is governed not only by feeding but also by other unknown factors

Efficiency and significance of multiple vocal signals in sibling competition

Animals can compete for resources by displaying various acoustic signals that may differentially affect the outcome of competition. We propose the hypothesis that the most efficient signal to deter opponents should be the one that most honestly reveals motivation to compete. We tested this hypothesis in the barn owl (Tyto alba) in which nestlings produce more calls of longer duration than siblings to compete for priority access to the indivisible prey item their parents will deliver next. Because nestlings increase call rate to a larger extent than call duration when they become hungrier, call rate would signal more accurately hunger level. This leads us to propose three predictions. First, a high number of calls should be more efficient in deterring siblings to compete than long calls. Second, the rate at which an individual calls should be more sensitive to variation in the intensity of the sibling vocal competition than the duration of its calls. Third, call rate should influence competitors' vocalization for a longer period of time than call duration. To test these three predictions we performed playback experiments by broadcasting to singleton nestlings calls of varying durations and at different rates. According to the first prediction, singleton nestlings became less vocal to a larger extent when we broadcasted more calls compared to longer calls. In line with the second prediction, nestlings reduced vocalization rate to a larger extent than call duration when we broadcasted more or longer calls. Finally, call rate had a longer influence on opponent's vocal behavior than call duration. Young animals thus actively and differentially use multiple signaling components to compete with their siblings over parental resources