Barn owl (Tyto alba) siblings vocally negotiate resources

Current theory proposes that nestlings beg to signal hunger level to parents honestly, or that siblings compete by escalating begging to attract the attention of parents. Although begging is assumed to be directed at parents, barn owl (Tyto alba) nestlings vocalize in the presence but also in the absence of the parents. Applying the theory of asymmetrical contests we experimentally tested three predictions of the novel hypothesis that in the absence of the parents siblings vocally settle contests over prey items to be delivered next by a parent. This 'sibling negotiation hypothesis' proposes that offspring use each others' begging vocalization as a source of information about their relative willingness to contest the next prey item delivered. In line with the hypothesis we found that (i) a nestling barn owl refrains from vocalization when a rival is more hungry, but (ii) escalates once the rival has been fed by a parent, and (iii) nestlings refrain from and escalate vocalization in experimentally enlarged and reduced broods, respectively. Thus, when parents are not at the nest a nestling vocally refrains when the value of the next delivered prey item will be higher for its nest-mates. These findings are the exact opposite of what current models predict for begging calls produced in the presence of the parents.     

Ambient noise increases missed detections in nestling birds

Ambient noise can mask acoustic cues, making their detection and discrimination difficult for receivers. This can result in two types of error: missed detections, when receivers fail to respond to the appropriate cues, and false alarms, when they respond to inappropriate cues. Nestling birds are error-prone, sometimes failing to beg when parents arrive with food (committing missed detections) or begging in response to stimuli other than a parent's arrival (committing false alarms). Here, we ask whether the frequency of these errors by nestling tree swallows (Tachycineta bicolor) increases in the presence of noise. We found that nestlings exposed to noise had more missed detections than their unexposed counterparts. We also found that false alarms remained low overall and did not differ significantly between noise and quiet treatments. Our results suggest that nestlings living in noisy environments may be less responsive to their parents than nestlings in quieter environments.     

Similarity in the begging calls of nestling Red‐winged Blackbirds

ABSTRACT Although individually distinct begging calls may permit parents to recognize their offspring, birds nesting in dense breeding colonies where fledglings intermingle might benefit from additional adaptations. For example, if the calls of all nestlings in a brood were similar, parents would need to recognize only one brood call instead of the identity calls of each nestling. We recorded nestling Red‐winged Blackbirds (Agelaius phoeniceus) to determine whether their calls function to identify individuals (identity call hypothesis) or broods (brood call hypothesis). We used spectrogram cross‐correlation and dynamic time warping as well as call duration, peak frequency, and frequency range to estimate the similarity of begging calls of nestling Red‐winged Blackbirds. We recorded individual nestlings on day 5 and on day 9 of the nestling period to determine whether calls of individuals were more similar than calls of different nestlings, and whether calls of broodmates were more similar than calls of nestlings from different broods. We found that calls of 8‐d‐old individuals were more similar than calls of different nestlings, but the calls of broodmates were not more similar than those of nestlings from different broods. These results were consistent with the identity call hypothesis. We then compared begging calls of pairs of nestlings recorded separately and together on day 9. We found that the calls of 8‐d‐old nestlings recorded together were more similar than when they were recorded separately. In addition, using playback of begging calls from normal broods and artificial “broods” constructed from the calls of single nestlings, we found that females returned with food sooner in response to the calls of single nestlings (with enhanced call similarity) than to those of normal broods. Our results suggest that similar begging calls may be beneficial for both nestlings and parents, with broodmates fed at higher rates when their calls are more similar and, after fledging, parents needing to recognize only one brood call instead of the identity calls of each fledgling.     

Begging in the absence of sibling competition in Wilson’s storm-petrels, Oceanites oceanicus

Begging displays of nestlings in multichick broods can signal both hunger and competitive ability. Studies of begging in species with single-chick broods exclude the influence of nestling competition and may provide especially useful models for the study of signalling during parent-offspring conflict. However, there is no evidence that chicks signal hunger by begging in the absence of sibling competition. I tested predictions of signalling models in a species with single-chick broods, the Wilson's storm-petrel. Chicks used two types of begging calls, ‘rhythmic’ calls and ‘long’ calls. I found that chicks conveyed information about their current body condition by begging. When their body condition was low, chicks increased the number and frequency of long begging calls, as well as the frequency of rhythmic calling. Parents responded to increased begging by regurgitating larger meals. The study thus demonstrates that the begging system can work in the absence of nestling competition. Chicks also called in the absence of their parents, but in this context they used only rhythmic calls and there was no correlation with current body condition. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Mouth colour components of begging are dynamic signals of quality in European starling nestlings

Offspring solicit food from their parents through begging signals. Nestling skin and flange coloration are begging signals that appear to convey information about nestling need or condition, and several experiments have shown that modifications of nestling coloration affect parental allocation decisions. However, it is important to examine the short‐term changes in these signalling components in response to food constraints since such dynamic changes are required for signals to indicate condition or need. Using a food deprivation experiment, we tested whether flange and skin reflectance in European starling Sturnus vulgaris nestlings change after a three‐hour interval. We investigated whether flange and skin reflectance changed according to the predictions arising from the ‘signal of quality’ or ‘signal of need’ hypotheses on the function of begging signals. We found that flange carotenoid and UV reflectance changed according to the signal of quality hypothesis with nestlings in good condition increasing their signal expression in response to the food deprivation, whereas those in poor condition decreased their signal expression. With the use of vision modelling, we show that changes in flange reflectance are detectable by starling parents. In contrast, we found a correlation going in the opposite direction for changes in skin UV reflectance. Nestlings with low lipid reserves increased their reflectance compared to nestlings with high reserves. However, vision modelling showed that short‐term changes in skin UV reflectance are not large enough to be detectable by the parents. Our study shows that flange carotenoid and UV reflectance are dynamic components of begging with short‐term variations that can be used by parents as signals of nestling quality.     

Begging in the absence of parents by nestling tree swallows

Begging by nestling passerine birds has become a model systemfor studies in animal communication. Although most beggingoccurs when parents arrive at the nest to feed (here called"primary begging"), it also occurs between feeding visits andimmediately after parents leave the nest. Begging in thesecontexts (here called "secondary begging") may have relativelylittle influence on the probability of receiving food, but could increase the overall cost of the signal and thus influence nestlingbegging strategies. The purpose of our study was to determinehow often tree swallow (Tachycineta bicolor) nestlings begin contexts other than to parents with food and to examinewhat factors influence the frequency of this begging. Secondarybegging ranged from 7% of measured begging responses at day2 to 30% by day 8 and was more frequent when the interval betweenparental feeding visits was relatively long and when the timeto respond to the arrival of parents with food was short. Increasesin both age and intervisit interval were associated with decreasesin nestling response times, suggesting that secondary beggingmay be related to the speed with which nestlings respond to stimuli. We discuss possible functions of secondary beggingand raise the possibility that it may, in fact, be an error.     

Great spotted cuckoos respond earlier to the arrival of feeding foster parents and perform less erroneous begging when hungry than their magpie host nest‐mates

In many bird species, parents usually feed the first nestling that starts to beg before its nest‐mates. The pressure to avoid missed feeds could trigger nestlings to perform in erroneous begging in absence of parents, which has the same costs as begging in the presence of parents but without any reward. So, nestlings should try to minimize both erroneous begging and missed feeds simultaneously. The threshold to start begging is predicted to be lower for hungry nestlings and for nestlings that are unrelated to their nest‐mates, because they suffer lower inclusive fitness costs when depriving nest‐mates of food. In line with this idea, we found that brood parasitic great spotted cuckoo nestlings responded sooner than their magpie nest‐mates when an adult arrived to the nest. Under laboratory conditions, nestlings of both species rarely incurred in erroneous begging when food was abundant, but under conditions of restricted food, magpie nestlings increased erroneous begging while cuckoo nestlings did not. Highly conspicuous begging in cuckoos results in an increased predation risk, which could have resulted in stronger selection pressures on cuckoos to avoid erroneous begging, probably resulting in better developed perceptual abilities, allowing cuckoos to perform better than their host nest‐mates.     

Parental Sex Differences in Food Allocation to Junior Brood Members as Mediated by Prey Size

Individual offspring within a brood may receive different amounts of provisioning from the male and female parents. Some hypotheses suggest that this bias is the result of an active and adaptive choice by parents. An alternative hypothesis is that feeding biases arise as a result of a constraint of fitting large prey items into small gapes. In an experiment with pied flycatchers, Ficedula hypoleuca , we tested for sex-biased allocation to junior nestlings in asynchronous broods and whether this could be explained by active parental choice or by passive allocation according to prey size and gape size. In both control broods and broods with experimentally increased degree of asynchrony, prey types did not differ between parents but females brought smaller prey than males at younger but not older nestling stages. At younger but not older nestling stages, the majority of feeds to junior nestlings were from females, and the smaller nestlings consumed smaller prey than older siblings. However, there was no evidence of active preference of small nestlings by females as parents did not differ in the tendency to bypass a begging senior nestling in order to feed a junior nestling. Provisioning rates by females were lower than those by males when nestlings were young and we suggest that foraging time constraints caused by the need to brood offspring result in females bringing smaller prey than males. In turn, the larger prey brought by males was more often transferred to larger offspring after the smaller ones failed to swallow it. In such cases, 'preferential' feeding of small nestlings by females may simply be a passive side effect of foraging constraints and gape-size limitations.     

Hatching Asynchrony Decreases the Magnitude of Parental Care in Domesticated Zebra Finches: Empirical Support for the Peak Load Reduction Hypothesis

Parent–offspring conflict over the supply of parental care results in offspring attempting to exert control using begging behaviours and parents attempting to exert control by manipulating brood sizes and hatching patterns. The peak load reduction hypothesis proposes that parents can exert control via hatching asynchrony, as the level of competition amongst siblings is determined by their age differences and not by their growth rates. Theoretically, this benefits the parents by reducing both the peak load of the offspring's demand and their overall demand for food and benefits the offspring by reducing the amplification of their competition. However, the peak load reduction hypothesis has only received mixed support. Here, we describe an experiment where we manipulated the hatching patterns of domesticated zebra finch Taeniopygia guttata broods and quantified patterns of nestling begging and parental feeding effort. There was no difference in the begging intensity of nestlings raised in asynchronous or experimentally synchronous broods, yet parental feeding effort was lower when provisioning asynchronous broods and particularly so when levels of nestling begging were low. Further, both parents acted in unison, as there was no evidence of parentally biased favouritism in relation to hatching pattern. Therefore, our study provided empirical support for the prediction that hatching asynchrony reduces the feeding effort of parents, thereby providing empirical support for the peak load reduction hypothesis.     

Group provisioning limits sharing conflict among nestlings in joint-nesting Taiwan yuhinas

Offspring often compete over limited available resources. Such sibling competition may be detrimental to parents both because it entails wasted expenditure and because it allows stronger offspring to obtain a disproportionate share of resources. We studied nestling conflict over food and its resolution in a joint-nesting species of bird, the Taiwan yuhina (Yuhina brunneiceps). We show that adult yuhinas coordinate their feeding visits, and that this coordination limits competition among nestlings, leading to a ‘fairer’ division of resources. Transponder identification and video-recording systems were used to observe adult feeding and nestling begging behaviours. We found that: (i) yuhinas feed nestlings more often in large parties than in small parties; (ii) feeding events occurred non-randomly in bouts of very short intervals; and (iii) food distribution among nestlings was more evenly distributed, and fewer nestlings begged, during large-party feeding bouts compared with small-party feeding bouts. To our knowledge, this is the first study in a cooperative breeding species showing that adults can influence food allocation and competition among nestlings by coordinating their feeding visits. Our results confirm the hypothesis that the monopolizability of food affects the intensity of sibling competition, and highlight the importance of understanding the temporal strategies of food delivery.     

Parental allocation of food to nestling tree swallows: the influence of nestling behaviour, sex and paternity

Parents are expected to invest more in young that provide the greatest fitness returns. The cues that parents use to allocate resources between their offspring have received much recent attention. In birds, parents may use begging intensity, position in the nest or nestling size as cues to provision the most competitive young or those most likely to survive. It may also benefit parents to invest in young differentially by sex or relatedness if the fitness returns of sons and daughters differ or broods are sired by multiple males. We examined the allocation of food to tree swallow, Tachycineta bicolor, nestlings in relation to their begging behaviour, size, sex and paternity. Provisioning by parents was not related to nestling size, sex or paternity. The begging behaviour of nestlings did not differ with respect to sex or paternity. Both parents were more likely to feed nestlings that begged first or were closer to the nest entrance, suggesting that parents allocate food resources in response to cues that nestlings control. As a consequence, brood reduction was facilitated by biased provisioning within the brood in addition to the nestling size hierarchies created by hatching asynchrony. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Barn swallow (<Emphasis Type="Italic">Hirundo rustica</Emphasis>) parents use past information to decide on food provisioning to the brood,but not to decide on allocation within the brood

The begging behavior of birds is thought to influence the allocation of food within the brood (allocation function) or to increase the total amount of food delivered to the brood (delivery function). Considering the nature of the feeding activity of passerines, in which hundreds of feeding events occur in a single day, parents may not necessarily decide the amount of resources allocated to each nestling/brood during a single feeding event. To examine this possibility, the relationship between begging intensity and parental responses to allocation and delivery functions was tested and modeled at multi-temporal scales in the barn swallow (Hirundo rustica). Comparison of models revealed that barn swallow parents differed in the temporal scales at which they respond to nestlings with respect to these two functions. While barn swallow parents decide which nestling to feed at a focal feeding event according to chick begging of the focal feeding event, they integrated past information on total begging effort to determine delivery rates to their offspring during the 14 and 6 min prior to the focal feeding event in males and females, respectively. These findings highlight that it is important to investigate parental response to begging at appropriate temporal scales when analyzing begging functions.     

Effect of alarm calling by male Red‐winged Blackbirds on nestling begging and female provisioning behavior

ABSTRACT Nestling begging and parental provisioning can attract nest predators and reduce reproductive success, so parents and their offspring might be expected to respond adaptively by minimizing predator‐attracting cues when predators threaten nests. Male Red‐winged Blackbirds (Agelaius phoeniceus) are well known for their antipredator alarm calls that contain information about the approach of potential nest predators. We examined the begging behavior of nestlings and the provisioning behavior of females in response to antipredator alarm calls of males to test the adaptive response hypothesis. Playback experiments provided no evidence that alarm calls function to switch off vocal begging; nestlings were equally likely to beg vocally during playback and control periods. Video recordings showed that male alarm calling had no significant effect on inappropriate vocal begging (in the absence of an adult), but significantly reduced the incidence of spontaneous calling (in the absence of begging). Adult females responded to male antipredator alarm calls by delaying their provisioning visits. In addition, although having no significant effect on use of nest‐arriving calls by females, male alarm calling significantly reduced their use of nest‐leaving calls. We conclude that nestling and female Red‐winged Blackbirds respond to male alarm calls in ways that might reduce the risk of predation, but nestlings beg vocally when females arrive to feed them, regardless of male alarm calling, perhaps to avoid a competitive disadvantage with broodmates.     

Nestling colouration is adjusted to parent visual performance in altricial birds

Hitherto, most of the investigation on the perceptual efficacy of begging signals has dwelled on how patterns of nestling colouration adjust to predominant nest luminosity. However, visual sensitivity of birds varies across species, which raises the question of whether colouration of traits involved in begging displays is adjusted to parent visual capacities. Here, by comparing nestling colouration and visual sensitivity across 22 altricial bird species, we provide a first test of this hypothesis. Firstly, we assessed differences in performance of typical UV‐tuned and violet‐tuned bird eyes when looking at the nestling traits under the light regimes prevailing at their nests. Secondly, while controlling for common ancestry in a comparative approach, we explored variation in colouration of nestlings in relation to parent visual system. The colour discrimination model indicated a general higher performance of the ultraviolet over the violet eye at detecting gape and body skin traits in either open‐ or hole‐nest light conditions. Gape colouration was associated with parental visual system as the nestlings of UVS species displayed more yellow and less pure ultraviolet mouths than the nestlings of VS species. Thus, our results agree with an adaptive parent–offspring communication scenario where the nestlings’ colours tuned the perception capacities of their parents.     

Nestling birds put their best flange forward

The offspring of caring parents may evolve specialized traits uniquely adaptive during their dependence on parental care. For example, the mouths of passerine nestlings are often bordered by enlarged and colorful rictal flanges expressed only during the nestling period. Although these traits are commonly hypothesized to act as visual signals during begging, non‐communication functions for the specialized mouth have been proposed as well. To test the hypothesis that nestling flange colors have evolved largely or exclusively as visual signals, I compared the reflectance of flange tissue that would be visible to parents during begging to that of flange tissue not exposed during begging in nestling house sparrows Passer domesticus and cliff swallows Petrochelidon pyrrhonota. Specifically, I tested the prediction that both condition‐dependent color parameters and those associated with visual conspicuousness would be expressed more intensely in tissue displayed during begging. Consistent with this prediction, flange tissue exposed during begging was brighter (reflected more total light), more UV‐rich, and had more intense carotenoid‐based coloration than hidden tissue. These differences do not exclude a non‐signaling function for flanges, but are consistent with the hypothesis that flange colors have evolved as visual signals.     

Parental alarm calls suppress nestling vocalization

Evolutionary models suggest that the cost of a signal can ensure its honesty. Empirical studies of nestling begging imply that predator attraction can impose such a cost. However, parents might reduce or abolish this cost by warning young of the presence of danger. We tested, in a controlled field playback experiment, whether alarm calls cause 5-, 8- and 11-day-old nestlings of the white-browed scrubwren, Sericornis frontalis, to suppress vocalization. In this species, nestlings vocalize when parents visit the nest ('begging') and when they are absent ('non-begging'), so we measured effects on both types of vocalization. Playback of parental alarm calls suppressed non-begging vocalization almost completely but only slightly reduced begging calls during a playback of parental feeding calls that followed. The reaction of nestlings was largely independent of age. Our results suggest two reasons why experiments ignoring the role of parents probably overestimate the real cost of nestling vocalizations. Parents can warn young from a distance about the presence of danger and so suppress non-begging vocalizations that might otherwise be overheard, and a parent's presence at the nest presumably indicates when it is safe to beg.     

Acoustic signalling of hunger and thermal state by nestling tree swallows

The begging displays used by altricial nestling birds to solicit care from parents include vigorous movements and loud calling. These begging signals have attracted considerable interest, mainly because their intensity seems excessive for the function of transmitting information about nestling need to parents. However, how information on need is encoded in the various components of the signal, especially its acoustic components, is poorly understood. We examined how begging calls of large and small nestling tree swallows, Tachycineta bicolor, changed during a short period of food deprivation and cooling, as a first step in determining the role that various call characteristics played in advertising nestling need. In contrast to previous studies, we examined several call variables, and related them not only to need for food but also need for warmth. When nestlings were deprived of food, their calls increased in rate and length. Large nestlings also increased the amplitude of their calls. When nestlings were cooled during food deprivation, they decreased the frequency of their calls and their call rate. The latter trend was especially evident in small nestlings. Our results suggest that begging calls carry information not only on the overall hunger level of broods, as emphasized in previous studies, but also on the size, hunger and thermal need of individual nestlings. Further tests are needed to determine whether parents use this information and whether begging calls are optimally designed to convey it. Copyright 2001 The Association for the Study of Animal Behaviour.     

杂色山雀亲代喂食与子代乞食间的行为

在育雏期,晚成鸟的子代一般都是由双亲共同来抚育,子代为了更好地存活,会用自己的方式竞争获得更多的食物和更好的生存空间,同时亲代也会根据子代的乞食信号来分配食物。2011年3～7月采用针孔摄像技术录制了杂色山雀(Parus varius)育雏期巢内亲代与子代间的行为,统计了亲鸟站位、雏鸟站位、雏鸟乞食强度及亲鸟的喂食情况等数据。分析结果表明:(1)雌雄亲鸟在巢中的站位各有特点,雄鸟在整个育雏期都喜欢站在距离巢口较近的位置;雌鸟站位不太固定,前期离巢口相对较远,中期和后期离巢口相对较近;(2)雏鸟离亲鸟越近,乞食强度越大,获得食物的机会就越多;离亲鸟越远的雏鸟越不爱乞食,所以站位对雏鸟的食物获得影响最大;(3)雌鸟承担主要的育雏任务,喂食频率远大于雄鸟;(4)育雏期的不同阶段雏鸟乞食强度、亲鸟喂食频率变化很大:中期雏鸟乞食强度最大,亲鸟喂食频率最高,后期雏鸟乞食强度最弱;(5)整个育雏期雌性亲本没有表现出明显的偏爱行为,但雄性亲本在中、后期更偏爱体型大的雏鸟。可见杂色山雀子代的行为和体型大小影响着亲代的食物分配,亲代也会根据雏鸟日龄调整站位和喂食行为。     

Facultative adjustment of pre‐fledging mass recession by nestling chimney swifts Chaetura pelagica

In species susceptible to mass‐dependent flight costs, mass recession prior to fledging may ensure that fledglings have appropriate wing loading. Our objectives were to determine if mass recession by chimney swift Chaetura pelagica nestlings is intrinsically controlled or facultatively adjusted by nestlings, and if mass recession is driven by changes in parental (i.e. reduced provisioning rates) or nestling (i.e. reduced begging) behavior. Nestling swifts (n = 50 in 17 broods) were divided into three treatment groups: controls, half‐weighted, or weighted. Half‐weighted and weighted nestlings had 0.6–0.7 or 1.2–1.3‐g lead weights, respectively, glued to body feathers on their backs during the period from 16 to 26 d post‐hatching. Weighted nestlings lost more mass than control and half‐weighted nestlings. After accounting for the added weights, control nestlings also had a higher wing loading than weighted nestlings. Video recordings revealed that provisioning rates of adult swifts did not vary throughout the nestling period, but the percent time nestlings spent begging increased slightly with age. Differences in mass recession among nestlings in different treatment groups resulted in convergence toward similar wing loading values likely optimal for flight efficiency. Mechanism(s) involved in this process remain unclear because provisioning rates were similar (from day 12 to 26 post‐hatching) whereas percent begging time by nestlings tended to increase with nestling age. However, weighted nestlings may have lost more mass than control nestlings by soliciting less food from adults than siblings, being more active, losing more water due to tissue maturation, or through some combination of two or more of these factors.     

Complex feeding behaviour by magpies in nests with great spotted cuckoo nestlings

Parent decisions about food allocation are usually based on simple time‐saving rules that optimize their own fitness; however, they can sometimes vary depending on the prevailing ecological conditions both outside and inside the nest. Parent–offspring interactions also become more complex when parents suffer from brood parasitism, which implies that they care for the parasite's eggs and unrelated young. The great spotted cuckoo Clamator glandarius is a specialist brood parasite that uses the magpie Pica pica as its primary host. Here, by filming food allocation by magpie parents in natural non‐parasitized and experimentally parasitized and non‐parasitized magpie nests, we have found that magpie provisioning behaviour is highly complex including two types of feedings apart from normal ones. First, false feedings, when the parent touched the chick's beak but did not leave any food, occurred more frequently when feeding a cuckoo than when feeding magpie nestlings. Second, two types of what we have called coax feedings: 2a) when magpie parents induce a nestling to beg by waking it up by touching it softly with the beak, and 2b) when parents disregard begging signals (always from brood parasitic great spotted cuckoos) while coaxing one non‐begging nestling (always one of their own) to feed it. We suggest that brood parasitism, involving selfish excessively begging nestlings, could have acted as a selective pressure for both false and coax feedings to evolve, as both imply ignoring nestlings that beg too much. We also discuss that these parental responses could have evolved either by a discrimination without recognition mechanism, or, more probably, by a recognition‐based discrimination mechanism.