Parent-offspring recognition in bank swallows (Riparia riparia): II. Development and acoustic basis

In study 1, bank swallow (Riparia riparia) chicks were exchanged with like-aged chicks from other broods. Parents accepted chicks that were transferred into their nests at age 15 days or younger; rejection began to occur at 16 to 17 days. In study 2, chicks' vocalizations were recorded in the burrow. We found that an immature begging call given by young chicks is replaced by a ‘signature’ call at 15 to 17 days of age. An acoustic analysis suggested that these calls are individually distinctive. Study 3 was a playback experiment designed to test whether the chicks' signature calls are a sufficient cue for parental recognition. We found that parents would approach a speaker broadcasting the calls of their chicks in preference to one simultaneously broadcasting the calls of alien chicks. The pattern of results suggests that parental recognition is based on the chicks' signature calls and that development of recognition is dependent on the development of the call.     

Individual recognition of parental calls by bank swallow chicks (Riparia riparia)

Antiphonal responses of newly fledged bank swallows (Riparia riparia) to the calls of their parents during the joined flights to and from the home burrow suggest that recognition of parents by offspring is a mechanism promoting family cohesion, a necessary condition for the adequate provisioning of chicks in this highly colonial species. Paternal and maternal calls used while leading chicks revealed higher inter-than intra-individual variability in temporal and frequency parameters, indicating that these calls could reliably identify individuals. When tested in the field at the onset of the fledging period, chicks answered significantly more playbacks of parent calls than of alien adult calls. Quantified observations of the rearing activity of the parents suggest that chick-parent recognition is based on familiarity, because of the chicks' differential exposure to the parental calls prior to fledging. During the development of the chicks in the burrow, both parents (which share the rearing activities equally) often call before feeding the chicks.     

Acoustic parameters of begging calls indicate chick body condition in Wilson’s storm-petrels <Emphasis Type="Italic">Oceanites oceanicus</Emphasis>

Begging behaviour as a key element in the parent–offspring conflict has been studied in many avian species. These types of studies have nearly exclusively been based on call counts, and it is still not entirely clear whether begging calls themselves contain any information. We studied begging behaviour in Wilson’s storm-petrel Oceanites oceanicus, a small procellariiform seabird. This species provides the opportunity to study the signalling value of begging calls in the absence of potentially confounding factors such as nestling competition, previous feeding experiences and predation pressure. We applied a new method using a semi-automatic spectrogram analysis software that measures the acoustic parameters of begging calls. Our analysis revealed that the frequency parameters of begging calls reflect chicks’ current body condition, with chicks in poorer condition uttering calls at higher frequencies. Chicks uttering higher pitched calls also received larger meals. Our study shows that certain acoustic parameters of begging calls can indicate the state of a chick in Wilson’s storm-petrels.     

Stimulus features of chicks and other factors evoking parental protective behaviour in ring-billed gulls

We examined the role of certain stimuli in parental approach and protection of ‘straying’ chicks by noting the response of ring-billed gulls, Larus delawarensis, to different stimuli placed within their territories. Playback of chick calls and presentation of surgically devocallzed chickes each elicited and oriented parental approach and protective behaviour, although a combination of the two was more effective. Other factors affecting a parent's reaction included the parent's reproductive state whether other offspring were present, the chick's location, and the chick'ssize relative to the parent's own chicks. These response tendencies should increase a gull's probability of responding selectively toward its own chicks before it can individually recognize them, while rejecting foreign chicks.     

Siberian crane chick calls reflect their thermal state

Chicks can convey information about their needs with calls. But it is still unknown if there are any universal need indicators in chick vocalizations. Previous studies have shown that in some species vocal activity and/or temporal-frequency variables of calls are related to the chick state, whereas other studies did not confirm it. Here, we tested experimentally whether vocal activity and temporal-frequency variables of calls change with cooling. We studied 10 human-raised Siberian crane (Grus leucogeranus) chicks at 3–15 days of age. We found that the cooled chicks produced calls higher in fundamental frequency and power variables, longer in duration and at a higher calling rate than in the control chicks. However, we did not find significant changes in level of entropy and occurrence of non-linear phenomena in chick calls recorded during the experimental cooling. We suggest that the level of vocal activity is a universal indicator of need for warmth in precocial and semi-precocial birds (e.g. cranes), but not in altricial ones. We also assume that coding of needs via temporal-frequency variables of calls is typical in species whose adults could not confuse their chicks with other chicks. Siberian cranes stay on separate territories during their breeding season, so parents do not need to check individuality of their offspring in the home area. In this case, all call characteristics are available for other purposes and serve to communicate chicks’ vital needs.     

Parent–offspring recognition in the Brown-headed Parrot Poicephalus cryptoxanthus

Recognition by vocal characteristics between parents and their offspring is thought to be ubiquitous in colonially nesting avian species. The Brown-headed Parrot Poicephalus cryptoxanthus nests in hollows in trees. However, when the chicks fledge they leave the nest and for the following three weeks spend their time in a tree where they are fed by their parents. As the fledglings are mobile and cryptic, returning parents must locate their own chicks. In this study a series of playback experiments was carried out, which showed that the chicks recognise their parents by voice. A simultaneous mirror experiment indicated that recognition was not reciprocated, although there may be alternative explanations for this behaviour. When the young began to forage with their parents, vocal stimuli did not induce any response from adult or chick. We suggest that vocal recognition becomes secondary to visual recognition as development proceeds.     

The role of parent–offspring interactions during and after fledging in the Black-legged Kittiwake

Most bird species endure a high mortality at fledging, and selection should favour parental behaviour diminishing these costs. Post-fledging parental care varies greatly among species and is often linked to parent–offspring recognition. In the Black-legged Kittiwake (Rissa tridactyla), fledglings need to return to the natal nest to be fed by their parents until independence. Rejections of fledglings by non-parent adults may be fairly violent, and parents are expected to recognize and help their chicks at the time of first return. However, previous cross-fostering experiments pointed out that parents are not able to recognize their chicks up to 15 days before fledging. In this paper, we study the behaviour of both parents and juveniles at fledging. We found that parents answered significantly more to their fledgling's calls than to those of others. Compared to silent juveniles, juveniles that called before landing were more likely to be accepted by their parents. No such pattern was observed with foreign juveniles, indicating that fledglings’ voice may carry individual identity. Furthermore, fledglings found their way back to the natal nest faster when parents attended the natal nest and reacted to their offspring's calls than when they were absent or inactive. Such interactions may therefore diminish juvenile mortality at fledging.     

Parental and chick responses of Laughing Gulls (Leucophaeus atricilla) to increase of flight costs and brood size

One main line of thought in life history theory is that investment in offspring must be balanced to minimize negative impacts on adult survival and future breeding. Seabirds have been regarded as fixed investors, although they exhibit a whole gradient of life history traits. We studied the consequences of brood size (one and three chicks) and of increased flight costs to one mate of a pair (3- and 5-cm trimming of the edge of the primary feathers) on parental response and on survival and body condition of chicks of Laughing Gulls (Leucophaeus atricilla). Parent gulls modified their nest attendance when their mate was handicapped, in a pattern dependent on the sex of the latter. Trimming of males affected chicks more severely than that of females. On its part, brood size affected amount of feeding sessions. Both chick body condition and survival were negatively affected by larger broods and by increased flight costs of one of their parents, especially when it was the male. Overall, parental inversion exhibited adjustments depending on the requirements of the brood and the fact that males compensated better the increased flying costs of their mates than vice versa. Despite a certain capacity by the males to compensate for the increased flight costs of the females, compensation was insufficient, and much less so in females, especially in larger broods, affecting chicks’ body condition and survival.     

Reliable Information Content and Ontogenetic Shift in Begging Calls of Grey Warbler Nestlings

The most critical assumption of communication models regarding parent–offspring conflict is that food solicitation displays of genetic offspring are honest signals to elicit beneficial parental care. A critical requirement of honesty is the reliable change of perceivable aspects of begging calls with physiological needs. We experimentally tested whether and how the acoustic structure and begging call rate of individual Grey Warbler Gerygone igata nestlings change with hunger level and age. We also examined a rarely documented component of chick begging calls, namely the temporal dynamics of acoustic modulation after nestlings heard parental feeding calls. Begging call structure narrowed in frequency range and, surprisingly, decreased in amplitude as chick hunger levels increased. We also found that begging calls changed with chick age, with the frequency increasing and the duration decreasing for older chicks. These results indicate that the acoustic properties of nestling Grey Warbler begging calls are complex and may be used to signal several aspects of nestling traits, including hunger level and age (or size, a correlate of age). Overall, begging calls of Grey Warbler chicks appear to be honest, implying that parents are likely to benefit from relying on the acoustic features of their progeny’s calls which predict chick need. Our results have important implications regarding the reliability and information content of nestling solicitation signals for the brood parasite shining cuckoo Chrysococcyx lucidus exploiting Grey Warbler parental care, in that these begging‐call mimetic specialist cuckoos might also need to match closely the dynamics of acoustic features of their host chicks’ calls.     

Relationships between brood size and parental provisioning performance in chinstrap penguins during the chick guard phase

 Chinstrap penguins (Pygoscelis antarctica) normally lay two eggs, but brood size is often reduced by mortality during incubation or after hatching. We hypothesized that this variation in brood size would affect the parents’ foraging behavior and their chick provisioning performance. We studied patterns of adult foraging trip duration and frequency, food load delivery, and chick growth rates in relation to brood size during the guard phase in four breeding seasons (1991–1994) on Seal Island, Antarctica. Within a given year, parents with two chicks made more frequent foraging trips to sea and may have transported larger food loads to the nest; however, the duration of foraging trips was unrelated to brood size. Overall, parents with two chicks spent ∼15% more time at sea than parents with only one chick. Both the frequency and duration of foraging trips varied between years. Foraging trip duration may partly reflect the birds’ foraging radius, which probably varies with time in response to shifts in krill distribution. Chick growth rate varied betwen years, but was related to brood size only in 1992, when chicks from two-chick broods grew significantly more slowly than chicks from one-chick broods. Food loads transported to chicks, as well as chick growth rates, were highest in 1994, when concurrent hydroacoustic studies indicated that regional krill biomass was severely depressed. This apparent anomaly suggests that the spatial scale of the krill survey may have been too coarse to detect some high-density krill aggregations within the penguins’ foraging range. Received: 26 September 1995 / Accepted: 12 May 1996     

On the responses of laughing gull chicks (Larus atricilla) to the calls of adults I. Recognition of the voices of the parents

Twelve matched pairs of parent-raised and hand-raised laughing gull chicks from the same clutches were tested, at 6 to 8 days post-hatching, with successive presentations of playback of a recording of calls of their parents and recording of calls of parents from a neighbouring nest, in a situation in which they could approach or withdraw from the sound.The parent-raised chicks oriented towards the sound, moved towards the sound and called when the calls of their own parents were played; they tended to withdraw from the sound and sit silently when the calls of the neighbouring parents were played.The hand-raised chicks fled from the sound and crouched in silence in response to both calls of their parents and the calls of neighbours. They were more vocal and active in periods without sound than were the parent-raised chicks.It is concluded that laughing gull chicks in nature learn to recognize the individual characteristics of the calls of their parents, and to react positively to the calls, through interaction with their parents prior to the age of 6 days post-hatching.     

Feeding frequency influences crèching age in the Dalmatian pelican, Pelecanus crispus

Crèching behaviour is common in colonial seabirds; nevertheless, the factors inducing chicks to aggregate remain relatively poorly understood. It has been proposed that brood size, laying date and nest attendance are important factors in the formation of a crèche. Moreover, in most species of pelicans, chicks join crèches following the development of homoeothermy and coincident with the end of the brooding behaviour. We studied effects of feeding rate, nest attendance, brood size, laying date and homoeothermy on the age at which chicks entered the crèche at a colony of Dalmatian pelicans (Pelecanus crispus), in Srebarna, Bulgaria. Single chicks were fed more frequently than chicks from two-chick broods. Unlike American white pelicans (Pelecanus erythrorhynchos), Dalmatian pelicans maintained brooding behaviour a further 9 days after chicks had developed thermoregulation abilities. In contrast to nests with two chicks, nests with only one chick were never left unattended by the parents before the chick reached the crèching stage. Laying date, nest attendance and brood size did not affect the age that the chick entered the crèche. The age the chick entered the crèche was not correlated with the age of homoeothermy acquisition, but chicks significantly joined the crèche at younger ages when the mean number of feeds per chick per day during the rearing period in the nest was higher. This result suggests an implication of growth rate in the crèching age. Joining the crèche earlier can provide benefits that could have strong implications for the chicks’ future reproductive lives.     

Cocktail-party effect in king penguin colonies

The king penguin, Aptenodytes patagonicus, breeds without a nest in colonies of several thousands of birds. To be fed, the chick must recognize the parents in a particularly noisy environment using only vocal cues. The call an adult makes when seeking the chick is emitted at a high amplitude level. Nevertheless, it is transmitted in a colonial context involving the noise generated by the colony and the screening effect of the bodies, both factors reducing the signal-to-noise ratio. In addition, the adult call is masked by a background noise with similar amplitude and spectral and temporal characteristics, enhancing the difficulty for the chick in finding its parents. We calculate that the maximum distance from the caller at which its signal can be differentiated from the background noise (signal-to-noise ratio equal to 1) should not exceed 8 to 9 m in a feeding area. But our tests show that, in fact, chicks can discriminate between the parental call and calls from other adults at a greater distance, even when call intensity is well below that of the noise of simultaneous calls produced by other adults. This capacity to perceive and extract the call of the parent from the ambient noise and particularly from the calls of other adults, termed the ''cocktail-party effect'' in speech intelligibility tests, enhances the chick''s ability to find its parents.     

Parent-offspring recognition in the barn swallow (Hirundo rustica)

Past research has shown that in the colonial bank swallow and cliff swallow, parents and offspring recognize one another by calls, enabling parents to locate and feed only their own chicks. To test the hypothesis that recognition traits are adaptations to coloniality, we examined parent-offspring recognition in the related but non-colonial species, the barn swallow. To allow cross-species comparisons, we used the same methods that had been used in the previous swallow studies. We carried out four studies. (1) Cross-fostering experiments done immediately before fledging gave no evidence of recognition. (2) In a playback experiment, parents gave no indication that they recognized the calls of their offspring. (3) In another playback experiment, chicks responded more strongly to the calls of their parents than to the calls of unrelated adults. The degree of recognition, however, was somewhat weaker than that seen in the two colonial swallow species. (4) Newly fledged young did not creche after fledging. Rather, family groups generally stayed apart from one another for almost 2 weeks, while parental care was still being given. The comparative data suggest that parent-offspring recognition via individually distinctive cues evolved in response to the intermingling of young associated with colonial living, and it thus weakly expressed in barn swallows compared to cliff swallows and bank swallows.     

Parental provisioning behaviour in a flock-living passerine, the Vinous-throated Parrotbill Paradoxornis webbianus

The amount of food delivered by parents to their chicks is affected by various life history traits as well as environmental and social factors, and this investment ultimately determines the current and future fitness of parents and their offspring. We studied parental provisioning behaviour in the Vinous-throated Parrotbill Paradoxornis webbianus, a species with an unusual social system that is characterised by flock-living, weak territoriality and variable nesting dispersion. Parental provisioning rate had a positive influence on chick mass gain, suggesting that provisioning rate is an effective measure of parental investment in this species. Males and females fed nestlings at approximately the same rate, and no other carers were observed at nests. Parents coordinated provisioning rates so that they mostly fed chicks synchronously. However, the extent to which parents coordinated provisioning was associated with their social environment, synchrony being positively related to local breeding density and negatively to nearest neighbour distance. The rate at which parents provisioned nestlings showed the same relationships with social measures, being greatest at higher density and when neighbours were closer. Visit rate was also related to chick age, but not to brood size, brood sex ratio, extra pair paternity, laying date, temperature, parents’ body characters, time of day or year. We conclude that a breeding pairs’ social environment plays an important role in determining parental investment, probably through its effects on the opportunities that parents have for foraging with conspecifics.     

Recognition of Young in A Colonially Nesting Bird

Parents ought to restrict costly parental care to their genetic offspring and, particularly when the risk of misdirecting care is high, parent‐offspring recognition may evolve. I tested whether adult cave swallows, which nest in dense colonies and feed fledglings in mixed‐family groups, discriminate against unrelated young, using temporary chick transfers at two nestling ages and a cross‐fostering experiment. Temporary chick transfers indicated that parents bias feedings toward their own offspring near fledging (18 d) but not at about halfway through the nesting period (10 d). I also examined how parents learn to identify their offspring by cross‐fostering young 3 d after hatching and testing parental response 2 wks later. Adults did not favor their own offspring over unrelated nestlings when both were unfamiliar to the focal parents. However, when parents encountered two of their own offspring, one of which was reared by foster parents, they preferentially fed the familiar nestling. By recognizing young, cave swallow parents reduce some risks of misdirected parental investment (mobile fledglings) but not others (extra‐pair young and intraspecific brood parasitism).     

Darwin's Finch Begging Intensity Does Not Honestly Signal Need in Parasitised Nests

Parental care should be selected to respond to honest cues that increase offspring survival. When offspring are parasitised, the parental food compensation hypothesis predicts that parents can provision extra food to compensate for energy loss due to parasitism. Chick begging behaviour is a possible mechanism to solicit increased feeding from attending parents. We experimentally manipulated parasite intensity from Philornis downsi in nests of Darwin's small ground finch (Geospiza fuliginosa) to test its effects on chick begging intensity and parental food provisioning. We used in‐nest video recordings of individually marked chicks to quantify nocturnal parasite feeding on chicks, subsequent diurnal chick begging intensity and parental feeding care. Our video analysis showed that one chick per brood had the highest parasite intensity during the night (supporting the tasty chick hypothesis) and weakest begging intensity during the day, which correlated with low parental care and rapid death. We observed sequential chick death on different days rather than total brood loss on a given day. Our within‐nest video images showed that (1) high nocturnal larval feeding correlated with low diurnal begging intensity and (2) parent birds ignored weakly begging chicks and provisioned strongly begging chicks. Excluding predation, all parasite‐free chicks survived (100% survival) and all parasitised chicks died in the nest (100% mortality). Weak begging intensity in parasitised chicks, which honestly signalled recent parasite attack, was not used as a cue for parental provisioning. Parents consistently responded to the strongest chick in both parasitised and parasite‐free nests.     

State-dependent parental care in the Antarctic petrel: responses to manipulated chick age during early chick rearing

Life histories are state-dependent, and an individual's reproductive decisions are determined by its available resources and the needs of its offspring. Here we test how a chick's needs for food and protection influence parental decisions in the Antarctic petrel, Thalassoica antarctica , where the parents, due to their long breeding lifespan, are expected to give priority to their own needs before those of the young. We exchanged one-day-old chicks with four-day-old chicks and studied how the parents subsequently provided care to the chick. The duration of the guarding period was adjusted, and parents left older chicks earlier and younger chicks later compared to controls. Three mechanisms were responsible for the adjustments. 1) Parents with an older chick co-ordinated fewer guarding spells whereas parents with a younger chick co-ordinated more guarding spells. 2) At the last guarding spell, i.e. where a parent left the chick alone before the partner returned, less time was spent with older chicks, and more time with younger chicks. 3) Foraging trip duration was shortened by parents given older chicks and prolonged by parents given younger chicks, probably in response to the chick's food demand. Hence, the parents responded quickly to the altered needs of the chick. Parents with high body mass guarded longer and were better able to co-ordinate the guarding spells compared to lighter parents. In conclusion, Antarctic petrels adjust reproductive decisions to their own, their mate's, and their chick's state, and they seem to respond to the chick's needs for both food and protection.     

A Cue-Isolation Experiment to Determine if Caspian Tern Parents Learn Their Offspring's Down Color

A ground-nesting gull or tern (Laridae) parent will attack a foreign conspecific chick that is substituted for the parent's chick at about the time the parent's chick is mobile. Presumably, parents discriminate among chicks using some combination of vocal, morphologic, and behavior cues. It is not known which cues are used. Morphologic variation in chicks' down potentially provides discrimination cues in some species of Laridae, such as the Caspian tern (Sterna caspia). To test discrimination based on the down color of chicks, a parent's chick was replaced sequentially with foreign conspecific chicks that were similar and dissimilar to the down color of the parent's chick. This cue-isolation experiment indicated that parents rejected dissimilar chicks more frequently. Thus, parents learn the down color of their chicks and then use this information as a basis for aggressive rejection of foreign chicks. A recognition system based on morphology allows parents to detect and reject foreign chicks with a probability greater than chance.     

Male chicks are more costly to rear than females in a monogamous seabird, the Common Murre

Feeding rates and mass loss during chick rearing were comparedfor individually marked parents of male and female Common Murre(Uria aalge) chicks at Great Island, Newfoundland, Canada, from1997–2001. Both parents in this socially monogamous seabirdspecies share parental care duties until colony departure, afterwhich the single chick is fed only by its father. Because murresprovision their single chicks with one clearly visible fishper trip, it is possible to accurately determine whether parentsdifferentially feed male and female chicks. Based on slightlygreater mass of males in adulthood, possibly favored by sexdifferences in breeding roles, we predicted that male nestlingswould be fed more than females. Fathers' feeding rate to sons,but not daughters, increased with chick age, whereas maternalfeeding rate increased with chick age for both sexes. When year-correctedfeeding rates of pairs rearing both sexes were compared, bothmothers and fathers fed their sons significantly more than theirdaughters in the later part of the chick-rearing period. Moreover,parents rearing male chicks lost mass at a significantly higherrate than those rearing females. There was no difference infledging age for sons and daughters. These results indicatethat differential parental allocation occurs and has measurablecosts even in a species with only slight adult sexual dimorphism.