Novel sources of (co)variation in nestling begging behavior and hunger at different biological levels of analysis

Biological hypotheses predicting patterns of offspring begging typically concern the covariance with hunger and/or development at specific hierarchical levels. For example, hunger drives within-individual patterns of begging, but begging also drives food intake among individuals within broods, and begging and food intake can covary positively or negatively among genotypes or broods. Testing biological phenomena that occur at multiple levels, therefore, requires the partitioning of covariance between traits of interest to ensure that each level-specific relationship is appropriately assessed. We performed a partial cross-fostering study on a wild population of great tits (Parus major), then used multivariate mixed models to partition variation and covariation in nestling begging effort and two metrics of nestling hunger within versus among individual nestlings and broods. At the within-individual level, we found that nestlings begged more intensely when hungrier (positive correlation between begging and hunger). However, among individuals, nestlings that were fed more frequently also begged more intensely on average (negative correlation between begging and hunger). Variation in nestling mass did not give rise to the negative correlation between begging and hunger among nestlings, but we did find that lighter nestlings begged more intensely than their heavier biological siblings, suggesting that this effect may be driven by a genetic component linked to offspring size. Our study illustrates how patterns of covariance can differ across biological levels of analysis and addresses biological mechanisms that could produce these previously obscured patterns.

Calling in the face of danger: Do nestling Red-winged Blackbirds (Agelaius phoeniceus) suppress begging in response to predator playbacks?

Nest predation is the most frequent cause of nest failure in birds such as the Red-winged Blackbird (Agelaius phoeniceus) that nest on or near the substrate. Nestlings should therefore exhibit adaptations to reduce the risk of nest predation. We tested the nestling antipredator hypothesis by examining the begging responses of Red-winged Blackbird nestlings to vocalizations of (1) an important nest predator (American Crow, Corvus brachyrhynchos), (2) a predator that rarely preys on nestlings (Cooper’s Hawk, Accipiter cooperii), and (3) a nonpredator (Northern Flicker, Colaptes auratus). We performed playbacks with (1) both parents present at the nest, (2) male at the nest, and (3) neither parent present. Following playback, we measured duration of nestling begging after the parent departed (begging persistence), bouts of otherwise normal begging when no parent was present (parent-absent begging), and calling without postural components of begging (nonpostural begging). When the male or both parents were present during playback, adults responded with alarm calls and nestlings significantly reduced parent-absent begging following American Crow and Cooper’s Hawk playbacks. Nonpostural begging was significantly reduced following Cooper’s Hawk playback, but there were no significant differences in the other begging variables. When neither parent was present, we found no significant differences in nonpostural begging in response to the 3 playback types, but parent-absent begging was significantly reduced following American Crow and Cooper’s Hawk playbacks when compared to Northern Flicker playbacks. These results show that nestlings suppress their vocal begging in response to calls of predators including Cooper’s Hawks even though they are not common nest predators.

Effects of manipulated levels of predation threat on parental provisioning and nestling begging

Parental provisioning behavior is a major determinant of offspring growth and survival, but high provisioning rates might come at the cost of increased predation threat. Parents should thus adjust provisioning activity according to current predation threat levels. Moreover, life-history theory predicts that response to predation threat should be correlated with investment in current reproduction. We experimentally manipulated perceived predation threat in free-living great tits (Parus major) by presenting parents with a nest predator model while monitoring different aspects of provisioning behavior and nestling begging. Experiments were conducted in 2 years differing greatly in ecological conditions, including food availability. We further quantified male territorial aggressiveness and male and female exploratory tendency. Parents adjusted provisioning according to current levels of threat in an apparently adaptive way. They delayed nest visits during periods of elevated perceived predation threat and subsequently compensated for lost feeding opportunities by increasing provisioning once the immediate threat had diminished. Nestling begging increased after elevated levels of predation threat, but returned to baseline levels by the end of the experiment, suggesting that parents had fully compensated for lost feeding opportunities. There was no evidence for a link between male exploration behavior or aggressiveness and provisioning behavior. In contrast, fast-exploring females provisioned at higher rates, but only in the year with poor environmental conditions, which might indicate a greater willingness to invest in current reproduction in general. Future work should assess whether these personality-related differences in delivery rates under harsher conditions came at a cost of reduced residual reproductive value.

Context-dependent strategies of food allocation among offspring in a facultative cooperative breeder

Natural selection should favor adoption of parental strategies that maximize fitness when allocating investment among offspring. In birds, begging displays often convey information of nestling need and quality, allowing parents to make adaptive food allocation decisions. We investigated how adults utilized cues likely to represent nestling competitive ability (begging position) and need (begging intensity) and a cue independent of nestling control (nestling sex) to distribute food among nestlings in a facultative cooperative breeder, the black-throated tit (Aegithalos concinnus). We found that parents reduced their efforts when helped, suggesting that parents of helped broods would have the potential to satisfy nestling needs more than unhelped parents. This suggestion was supported by the fact that nestling mass increased faster in helped than in unhelped nests. We found no effect of nestling sex on food allocation, but, as predicted, we found that adults responded differently to begging signals in relation to the presence of helpers and brood size. First, helped parents were more responsive to nestling begging intensity than parents without helpers. Second, female parents and helpers had a stronger preference for nestling begging position in large than in small broods. Third, the preference for nestling begging position was greater for unhelped than for helped female parents. These results provide evidence that carers adjust their preference for different offspring begging signals based on availability of food resources.

Nestling begging calls increase predation risk by corvids

Despite nest predation being the most common cause of breeding failure in open-nesting birds, we have little insight into the cues used by nest predators when they search for nests. So far we have assumed that nest-predating birds are visually oriented while mammal predators to a large extent use scent and auditory cues like nestling begging calls. To evaluate how important nestling begging calls are for corvid nest predators searching for nests, I used artificial nests, which made it possible to find the real costs of the begging without mitigation by parental and nestling behavior. I used paired artificial nests, one with and one without nestling begging call playback. Within 10 days, 62.9% of the nests were predated. The analyses showed that nests with begging calls suffered a significantly higher predation rate than nests without begging calls, especially when the nests were placed close to corvid nests. Moreover, nests with begging calls were predated significantly earlier than nests without begging calls. In artificial nest pairs with both nests predated but on different days, nests with begging calls were predated first. In nest pairs with only one predated nest, nests with begging calls were predated most often. This experiment shows that nestling begging calls imply a cost in terms of increased and earlier nest predation, and that corvids use nestling begging calls as a cue to find and depredate bird nests, challenging earlier expectations.

Effect of Variation in Nestling Hunger Levels on the Begging Behaviour of Nestlings and the Provisioning Behaviour of Adult American Kestrels

Little is known about how variation in nestling begging intensity influences the behaviour of adult raptors and how responses of adult males and females to such variation might differ. Our objective was to manipulate the begging intensity of nestling American Kestrels (Falco sparverius) and examine the responses of adults. We studied 12 pairs of American Kestrels nesting in nest boxes from 1 March to 1 July 2014 at the Blue Grass Army Depot, Madison County, Kentucky. Nest boxes were modified with a separate compartment for a camcorder to record nestling behaviour, and a second camcorder was placed outside the nests to monitor adult behaviour. To manipulate nestling hunger levels, 12 to 26-day-old nestlings in six nests were deprived of food for 24 h and those in the other six nests were fed until satiated. At each nest, we alternated control (no treatment) and treatment (fed or food deprived) days over a 4 day period to minimise the possible effect of nestling age on adult and nestling behaviour. Nestling begging intensity differed among treatments, with nestlings in food-deprived nests begging with greater intensity after food deprivation and those in fed-treatment nests begging with less intensity after being fed. Adult male and female American Kestrels provisioned nestlings at similar rates, with both sexes feeding nestlings at higher rates after food deprivation and at lower rates after fed treatments. Thus, the begging behaviour of nestling American Kestrels varied with hunger level, and adult American Kestrels responded by adjusting provisioning rates. Although the response of adults to nestling begging suggests that natural selection might favour ‘dishonest’ begging to obtain more food, the potential costs of ‘dishonest’ begging, such as attracting predators, reduced immunocompetence, and loss of indirect fitness benefits if such begging negatively impacts siblings and parents, may outweigh any possible benefit.

Male parental investment reflects the level of partner contributions and brood value in tree swallows

Biparental care presents an interesting case of cooperation and conflict between unrelated individuals. Several models have been proposed to explain how parents should respond to changes in each other’s parental care to maximize their own fitness, predicting no change, partial compensation, or matching effort as a response. Here, we present an experiment in tree swallows (Tachycineta bicolor) in which we increased the parental care of females by presenting them, but not their mates, with additional nestling begging calls using automated playbacks. We performed this experiment in two populations differing in future breeding opportunities and thus the intensity of conflict over current parental care. We found that in response to a temporary increase in female parental effort, males in the northern population with lower sexual conflict matched the increased effort, whereas males in the southern population did not. We also found that increases in parental care during playbacks were driven by the females (i.e., females initiated the increased effort and their mates followed them) in the northern population but not the southern population. These results support the idea that with incomplete information about the brood value and need, cues or signals from the partner might become important in coordinating parental care.

Parental influence on begging call structure in zebra finches ( Taeniopygia guttata ): evidence of early vocal plasticity

Begging calls are signals of need used by young birds to elicit care from adults. Different theoretical frameworks have been proposed to understand this parent–offspring communication. But relationships between parental response and begging intensity, or between begging characteristics and proxies of a young’s need remain puzzling. Few studies have considered the adjustment of nestling begging features to previous experience as a possible explanation of these discrepancies. In this study, we tested the effect of a heterospecific rearing environment on individual developmental trajectories of the acoustic structure of nestling begging calls. Fifty-two zebra finch chicks were fostered either to Bengalese finch or to zebra finch parents, and begging calls were recorded at several stages of nestling development. Acoustic analyses revealed that the development of the spectral features of the begging calls differed between experimental conditions: chicks reared by Bengalese finches produced higher pitched and less broadband begging calls than chicks reared by conspecific parents. Differences were stronger in males than females and were not explained by differences in growth rate. We conclude that nestling begging calls can be plastic in response to social interactions with parents.

Low cost audiovisual playback and recording triggered by radio frequency identification using Raspberry Pi

Carrying out playbacks of visual or audio stimuli to wild animals is a widely used experimental tool in behavioral ecology. In many cases, however, playback experiments are constrained by observer limitations such as the time observers can be present, or the accuracy of observation. These problems are particularly apparent when playbacks are triggered by specific events or are targeted to specific individuals. We developed a low-cost automated playback/recording system, using two field-deployable devices: radio-frequency identification (RFID) readers and Raspberry Pi micro-computers. This system detects a specific passive integrated transponder (PIT) tag attached to an individual, and subsequently plays back the stimuli, or records audio or visual information. To demonstrate the utility of this system, we tagged female and male tree swallows from two box-nesting populations with PIT tags and carried out playbacks of nestling begging calls every time females entered the nestbox over a six-hour period. We show that the RFID-Raspberry Pi system presents a versatile, low-cost, field-deployable system that can be adapted for many audio and visual playback purposes. The low cost and the small learning curve make this set-up a feasible system for use by field biologists.