Sibling competition and parental control: Patterns of begging in parrots

Begging and food allocation patterns are the outcome of complex and repeated interactions between parents and young. In most systems studied, food allocation is regulated by begging and scramble competition. In contrast, little is understood about how nestling solicitation behaviours will evolve in systems where parents engage in complex patterns of food allocation. Parrots appear to be an excellent group in which to examine the shifting balance between sibling competition and parental control. Studies to date have shown that levels of sibling competition within parrot broods are low, possibly in response to parental control over food distribution. I assess what is known about the function of nestling begging in parrots and evaluate why begging signals appear to function differently in this group.

No experimental evidence that sibling competition induces young to switch nests in the colonial Alpine swift, Apus melba

Adoption is frequent in colonial animals where opportunities for dependent young to receive care from nonbiological parents are high. The departure of dependent young from their original family to seek adoption in neighbouring families is thought to be induced by sibling competition for access to limited resources provided by poor-quality parents. We tested this hypothesis in the colonial Alpine swift by manipulating the number of young reared per brood, with the prediction that offspring from enlarged broods switch nests more frequently than those from reduced broods. Although nestling swifts hatch with little locomotor activity, from 20 days until their first flight at 50-70 days they frequently move out of their nests to seek adoption in neighbouring families. Although nestlings reared in experimentally enlarged broods were lighter and their body mass at day 20 after hatching was more variable than in nestlings reared in reduced broods, there was no difference between the two treatments in the frequency of nests switching and in the age when nestlings switched nests for the first time. However, consistent with other evidence that nest switching by nestling swifts evolved as a strategy to reduce ectoparasite load, young from broods with naturally high numbers of the ectoparasitic louse fly Crataerina melbae were more prone to switch nests. This shows that ectoparasitism rather than sibling competition is a key proximate factor promoting the evolution of nest switching in the colonial Alpine swift. (c) 2006 The Association for the Study of Animal Behaviour Published by Elsevier Ltd. All rights reserved.

The vigilance components of begging and sibling competition

The conflict between siblings over how parental resources are divided has promoted the evolution of specific behaviour to outcompete each other. Young animals look out for parents' arrival in order to start begging as quickly as possible, since a rapid begging reaction increases the likelihood of being fed before nestmates. If the young can physically intercept the parents, selection might be operating on the offspring ability to monitor parent arrival (vigilance towards parents) and any sudden modifications in siblings' behaviour (vigilance towards siblings). To investigate the adaptive value of nestling vigilance in the context of family interactions, we recorded which direction barn owl Tyto alba siblings were facing in 89 two-chick broods before the first parental feeding visit of the night. Nestlings were more vigilant towards nest entrance than expected by chance suggesting that vigilance towards parents is an important component of sibling competition. When positioned near the nest-box entrance where parents predictably deliver food, the younger individual (i.e. junior) looked more towards the entrance than its older sibling. Thus, when the likelihood of obtaining a food item is relatively high, juniors are more vigilant than seniors to detect the incoming parent. When positioned at the back of the nest, the senior looked relatively more frequently towards its sibling than the junior did in the same situation. This suggests that when the likelihood of obtaining a food item is relatively low, seniors are more vigilant than juniors to observe their sibling. Because vigilance was not related to hunger level and prey obtaining, we propose the hypothesis that vigilance towards parents and siblings only indirectly influences the outcome of sibling competition.

Sibling competition and the risk of falling out of the nest

In birds, sibling competition encompasses several activities, one of which is jostling for position, that is, competing for the location in the nest where parents predictably deliver food items. We hypothesized that nestlings that compete by jostling for position may fall out of the nest either accidentally or because siblings push each other to reduce brood size. This hypothesis predicts that in a competitive environment needy nestlings trade-off the benefit of being fed against the cost of falling out of the nest. As a first attempt to evaluate this hypothesis, we experimentally manipulated the number of young per brood in the colonial Alpine swift, Apus melba. Nestlings fell out of their colony more frequently when reared in enlarged than in reduced broods. Because brood size manipulation affects not only the number of young per nest but also their body condition, we analysed an extended data set to disentangle these two factors. This analysis showed that, independently of brood size, nestlings in poor condition and those reared in broods where sibling differed markedly in weight were more likely to disappear from the colony. Nestling disappearance also occurred predominantly in nests close to the colony entrances. Although nestling swifts can wander in the colony and become adopted in neighbouring nests, we found no evidence that wandering per se increased the risk of falling out of the colony. Our study therefore highlights a novel cost of scramble competition.

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.

Effect of sibling competition and male carotenoid supply on offspring condition and oxidative stress

Early developmental conditions have major implications for an individual's fitness. In species where offspring are born simultaneously, the level of sibling competition for food access is intense. In birds, high sibling competition may subject nestlings to decreased growth rate as a result of limited food and increased levels of oxidative stress through high metabolic activity induced by begging behaviors. We manipulated the level of sibling competition in a natural population of great tits and assessed the consequences for nestling body condition and resistance to oxidative stress. In a full factorial design, we both augmented brood size to increase sibling competition and supplemented the male parents with physiological doses of carotenoids thereby doubling the natural carotenoid intake, aiming at increasing the males' investment in current reproduction and thereby decreasing sibling competition. Nestling body mass was reduced by the brood enlargement and enhanced by the carotenoid supplementation of fathers. Nestling resistance to oxidative stress, measured as total antioxidant defenses in whole blood, was not influenced by the treatments. Because nestlings experience high metabolic activities, an absence of an effect of sibling competition on free radicals production seems unlikely. Nestling body mass decreased and resistance to oxidative stress tended to increase with initial brood size, and hence these correlational effects suggest a trade-off between morphological growth and development of the antioxidant system. However, the result of the experimental treatment did not support this trade-off hypothesis. Alternatively, it suggests that nestling developed compensatory mechanisms that were not detected by our antioxidant capacity measure.

Signaling of need, sibling competition, and the cost of honesty

Young birds and mammals frequently solicit food by means of extravagant and apparently costly begging displays. Much attention has been devoted to the idea that these displays are honest signals of need, and that their apparent cost serves to maintain their honesty. Recent analyses, however, have shown that the cost needed to maintain a fully informative, honest signal may often be so great that both offspring (signaler) and parent (receiver) would do better to refrain from communication. This apparently calls into question the relevance of the costly signaling hypothesis. Here, I show that this argument overlooks the impact of sibling competition. When multiple signalers must compete for the attention of a receiver (as is commonly the case in parent–offspring interactions), I show that (all other things being equal) individual equilibrium signal costs will typically be lower. The greater the number of competitors, the smaller the mean cost, though the maximum level of signal intensity employed by very needy signalers may actually increase with the number of competitors. At the same time, costs become increasingly sensitive to relatedness among signalers as opposed to relatedness between signalers and receivers. As a result of these trends, signaling proves profitable for signalers under a much wider range of conditions when there is competition (though it is still likely to be unprofitable for receivers).

The evolution of begging: Signaling and sibling competition

In many species, young solicit food from their parents, which respond by feeding them. Because of the difference in genetic make-up between parents and their offspring and the consequent conflict, this interaction is often studied as a paradigm for the evolution of communication. Existent theoretical models demonstrate that chick signaling and parent responding can be stable if solicitation is a costly signal. The marginal cost of producing stronger signals allows the system to converge to an equilibrium: young beg with intensity that reflects their need, and parents use this information to maximize their own inclusive fitness. However, we show that there is another equilibrium where chicks do not beg and parents’ provisioning effort is optimal with respect to the statistically probable distribution of chicks’ states. Expected fitness for parents and offspring at the nonsignaling equilibrium is higher than at the signaling equilibrium. Because nonsignaling is stable and it is likely to be the ancestral condition, we would like to know how natural systems evolved from nonsignaling to signaling. We suggest that begging may have evolved through direct sibling fighting before the establishment of a parental response, that is, that nonsignaling squabbling leads to signaling. In multiple-offspring broods, young following a condition-dependent strategy in the contest for resources provide information about their condition. Parents can use this information even though it is not an adaptation for communication, and evolution will lead the system to the signaling equilibrium. This interpretation implies that signaling evolved in multiple-offspring broods, but given that signaling is evolutionarily stable, it would also be favored in species which secondarily evolved single-chick broods.

Individual vocal signatures in barn owl nestlings: does individual recognition have an adaptive role in sibling vocal competition?

To compete over limited parental resources, young animals communicate with their parents and siblings by producing honest vocal signals of need. Components of begging calls that are sensitive to food deprivation may honestly signal need, whereas other components may be associated with individual-specific attributes that do not change with time such as identity, sex, absolute age and hierarchy. In a sib-sib communication system where barn owl (Tyto alba) nestlings vocally negotiate priority access to food resources, we show that calls have individual signatures that are used by nestlings to recognize which siblings are motivated to compete, even if most vocalization features vary with hunger level. Nestlings were more identifiable when food-deprived than food-satiated, suggesting that vocal identity is emphasized when the benefit of winning a vocal contest is higher. In broods where siblings interact iteratively, we speculate that individual-specific signature permits siblings to verify that the most vocal individual in the absence of parents is the one that indeed perceived the food brought by parents. Individual recognition may also allow nestlings to associate identity with individual-specific characteristics such as position in the within-brood dominance hierarchy. Calls indeed revealed age hierarchy and to a lower extent sex and absolute age. Using a cross-fostering experimental design, we show that most acoustic features were related to the nest of origin (but not the nest of rearing), suggesting a genetic or an early developmental effect on the ontogeny of vocal signatures. To conclude, our study suggests that sibling competition has promoted the evolution of vocal behaviours that signal not only hunger level but also intrinsic individual characteristics such as identity, family, sex and age.

Effects of hatching asynchrony on sibling negotiation, begging, jostling for position and within-brood food allocation in the barn owl Tyto alba

When siblings differ markedly in their need for food, they may benefit from signalling to each other their willingness to contest the next indivisible food item delivered by the parents. This sib-sib communication system, referred to as 'sibling negotiation', may allow them to adjust optimally to investment in begging. Using barn owl (Two alba) broods. I assessed the role of within-brood age hierarchy on sibling negotiation, and in turn on jostling for position where parents predictably deliver food (i.e. nest-box entrance), begging and within-brood food allocation. More specifically, I examined three predictions derived from a game-theoretical model of sibling negotiation where a senior and a junior sibling compete for food resources (Roulin, 2002a, Johnstone and Roulin, 2003): (1) begging effort invested by the senior sibling should be less sensitive to the junior sibling's negotiation than vice versa; (2) the junior should invest less effort in sibling negotiation than its senior sibling but a similar amount of effort in begging; and (3) within-brood food allocation should be directly related to begging but only indirectly to sibling negotiation. Two-chick broods were created and vocalization in the absence (negotiation signals directed to siblings) and presence (begging signals directed to parents) of parents was recorded. In support of the first prediction, juniors begged at a low cadence after their senior sibling negotiated intensely, probably because negotiation reflects prospective investment in begging and hence willingness to compete. In contrast, the begging of senior siblings was not sensitive to their junior sibling's negotiation. In contrast to the second prediction, juniors negotiated and begged more intensely than their senior sibling apparently because they were hungrier rather than younger. In line with the third prediction, juniors monopolized food delivered by their parents when their senior sibling begged at a low level. The begging cadence of both the junior and senior sibling, the junior's negotiation cadence, the difference in age between the two nest-mates and jostling for position were not associated with the likelihood of monopolizing food. In conclusion, sibling negotiation appears to influence begging behaviour, which, in turn, affects within-brood food allocation. Juniors may negotiate to challenge their senior siblings, and thereby determine whether seniors are less hungry before deciding to beg for food. In contrast, seniors may negotiate to deter juniors from begging.

Parental influence on sibling rivalry in great tit, Parus major, nests

Sibling and parente-offspring conflicts arise mainly over the amount and distribution of parental care, especially food. In altricial bird species where the young depend on parents for obtaining food, parents may control sibling competition by the choice of their respective provisioning locations. In great tits, the parents use fixed provisioning positions on the nest rim that are determined early in the breeding cycle and maintained until. edging. The two parents may choose positions that are close to each other, or far apart, and thereby increase or relax the pressure for optimal feeding positioning among nestlings. As an inspiration to this study we previously found that the two parents provide food from closer positions if the nest is infested by ectoparasites. Here, we tested the hypothesis that the parental choice of relative provisioning locations could be strategically used to control nestling competition. We forced parents to feed from either one or two provisioning locations and assessed the induced change in nestling movement, weight gain, and food distribution among siblings. We show that the angular distance between male and female locations influences the level of behavioural competition and affects nestling weight gain and food distribution. It is the first evidence for hole-nesting birds, where it was assumed that the nestling closest to the entrance hole was fed first, that the apparent choice of feeding positions by parents could be a way of controlling sibling competition and thereby also taking partial control over the outcome of parente-offspring conflict. (c) 2007 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Begging and food distribution in crimson rosella (Platycercus elegans) broods: why don't hungry chicks beg more?

Altricial nestlings solicit food by begging and engaging in scramble competition. Solicitation displays can thus signal both hunger and competitive ability. I examined nestling solicitation and parental responses in crimson rosellas (Platycercus elegans), a species in which parents engage in complex patterns of food allocation and appear to control the distribution of food. By manipulating the hunger of individual chicks and entire broods, I assessed how chick behaviours and parental food allocation varied with hatching rank, level of hunger, and intensity of nestling competition. Last-hatched chicks begged more than first-hatched chicks irrespective of individual hunger levels. The two parents combined fed individually hungry chicks more, but mothers and fathers varied in their responses to begging chicks: fathers fed last-hatched chicks in proportion to their begging intensity, whereas mothers fed chicks equally. Since fathers generally allocate more food to first-hatched chicks, fathers appear to use begging rates to adjust food allocation to non-preferred chicks within the brood. When I manipulated brood hunger levels, begging rates increased for first- and last-hatched chicks suggesting that chick begging rates are sensitive to the level of competition. This study shows that begging by rosella chicks does not correlate with hunger in a straightforward way and that the primary patterns of food allocation by parents art: not influenced by chick begging. Thus the benefits of increased begging may be limited for nestlings in this species.

Laying date and laying sequence influence the sex ratio of crimson rosella broods

We examine the patterns of sex allocation in crimson rosellas Platycercus elegans, a socially monogamous Australian parrot. Overall, 41.8% of nestlings were male, a significant female bias. However underlying this population-level bias were non-random patterns of sex allocation within broods. Broods produced early in the season were female-biased, but the proportion of males in a brood increased as the breeding season progressed. Female rosellas may obtain greater fitness benefits from early-fledging daughters than sons because daughters can breed as 1-year-olds whereas sons do not breed until they are at least 2 years old. Laying date and laying sequence also interacted to influence the sex ratio of eggs. The sex of early-laid eggs strongly followed the brood level pattern, whereas the sex of middle- and late-laid eggs did not change significantly as the season progressed. Nevertheless, late-laid eggs were very unlikely to be male at the end of the season. We argue these differing seasonal patterns reflect the relative costs and benefits to producing early-hatched males and females at different times of the season. Female rosellas appear to maximise the probability that daughters are able to breed early but to minimise competitive asymmetries within the brood. In particular, late-hatched male chicks are disadvantaged if their oldest sibling is male, explaining the dearth of broods containing late-hatched males at the end of the breeding season.

Effects of brood size manipulation and common origin on phenotype and telomere length in nestling collared flycatchers.

BACKGROUND: Evidence is accumulating that telomere length is a good predictor of life expectancy, especially early in life, thus calling for determining the factors that affect telomere length at this stage. Here, we investigated the relative influence of early growth conditions and origin (genetics and early maternal effects) on telomere length of collared flycatchers (Ficedula albicollis) at fledging. We experimentally transferred hatchlings among brood triplets to create reduced, control (i.e. unchanged final nestling number) and enlarged broods. RESULTS: Although our treatment significantly affected body mass at fledging, we found no evidence that increased sibling competition affected nestling tarsus length and telomere length. However, mixed models showed that brood triplets explained a significant part of the variance in body mass (18%) and telomere length (19%), but not tarsus length (13%), emphasizing that unmanipulated early environmental factors influenced telomere length. These models also revealed low, but significant, heritability of telomere length (h(2) = 0.09). For comparison, the heritability of nestling body mass and tarsus length was 0.36 and 0.39, respectively, which was in the range of previously published estimates for those two traits in this species. CONCLUSION: Those findings in a wild bird population demonstrate that telomere length at the end of the growth period is weakly, but significantly, determined by genetic and/or maternal factors taking place before hatching. However, we found no evidence that the brood size manipulation experiment, and by extension the early growth conditions, influenced nestling telomere length. The weak heritability of telomere length suggests a close association with fitness in natural populations.

Sib-sib communication and the risk of prey theft in the barn owl Tyto alba

Conflicts among siblings are widespread and their resolution involves complex physical and communication tools. Observations in the barn owl Tyto alba showed that siblings vocally communicate in the absence of parents to negotiate priority of access to the impending food resources that parents will bring. In the present paper, we hypothesize and provide correlative evidence that after a parent brought a food item to their progeny, sibling competition involves vocal sib-sib communication. A food item takes a long time to be entirely consumed, and hence siblings continue to compete over prey monopolization even after parents gave a food item to a single offspring. When physical competition is pronounced and thereby the risk of prey theft is high, the individual that received a prey item consumes it in a concealed place. Concomitantly, nestlings vocalize intensely probably to indicate their motivation to siblings to not share their food item, since this vocal behaviour was particularly frequent in younger individuals for which the risk of being robbed is higher than in their older siblings. Furthermore, nestlings consumed more rapidly a food item when their siblings vocalized intensely presumably because the intensity of siblings' vocalizations is associated with a risk of prey theft. Our correlative study suggests that sibling competition favoured the evolution of sib-sib communication under a wide range of situations.