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.     

Chick begging as a signal: are nestlings honest?

Begging by dependent avian offspring is known to correlate withhunger level, and parents use this as a signal of brood demandto adjust their chick feeding behavior. While there is informationon how each chick adjusts its begging to its own condition,little is known of how chicks adjust to the state of their nestmates. In two experiments we manipulated the competitive environmentof individual European starling (Sturnus vulgaris) chicks byaltering the state of nest mates while holding the state oftarget chicks constant In the first experiment we placed thetarget chick's nest mates in neighboring nests with brood sizesof two, five, or eight chicks. Following the manipulation wereturned them to their own nests and recorded begging behavioron videotape. In the second experiment we separated a targetchick from its siblings and manipulated feeding level in thelaboratory. The siblings were fed at one of three levels; meanwhile,all the target chicks were fed at the intermediate level. Afterthe manipulation we placed the target chicks with their siblingsand recorded their begging in response to an artificial stimulus.In neither experiment was the begging effort of the unmanipulatedtarget chicks affected by the changes in begging behavior oftheir siblings. This result supports the view that begging isa reliable signal of individual chick state and does not involveresponses to the effort of nest mates.     

Sibling competition and siblicide in asynchronously-hatching broods of the cattle egret Bubulcus ibis

Feeding behaviour and sibling competition were observed in nine families of the cattle egret (Bubulcus ibis) from blinds during 1359 nest-h throughout the nestling period. During days 0–19, size differences among siblings were clear; begging behaviour of chicks changed with time. At least one parent always attended the nest. Food boluses regurgitated early within a feeding period were received by senior chicks more often than by juniors. When any two siblings begged for food at the same time, the elder and younger received the first bolus on 65% and 35% of occasions respectively. Between days 20 and 39, the frequency of begging reached a peak. Begging behaviour became intense and stereotyped. The number of boluses received per begging declined rapidly, especially for junior chicks. In large broods, the success rate of begging was lower and fights occurred among siblings, especially among juniors. Out of 256 dyadic fights, the elder sibling won 85, lost one, and tied 171. The youngest chick died in two broods, apparently as the result of these fights (siblicide). No parents interfered in fights among their offspring. After day 40, the frequency of begging decreased gradually and ceased by day 80, No chicks died in the last period, although the frequency of fights in all large broods remained high.     

Prolactin and parental behavior in Adélie penguins: effects of absence from nest, incubation length, and nest failure

Adélie penguin (Pygoscelis adeliae) males and females, nesting in Antarctica, alternate attendance at the nest with absences of many days to forage at sea. We investigated the importance of tactile input from egg and chicks on prolactin levels by observing nest attendance patterns and obtaining blood samples (1) during the first nest exchange of the incubation stage, (2) from birds whose incubation period was artificially increased or decreased by about 10 days, and (3) from birds whose nests had failed. Prolactin levels in females after 8 to 11 days of absence from the breeding colony did not differ from those in incubating males and did not change after females resumed incubation. Moving eggs between nests resulted in nests in which chicks hatched after about 26, 36 (normal), or 46 days. Duration of incubation did not affect prolactin levels in the parents measured during incubation, at the pip stage, hatch stage, or early brood stage. Adults first left their chicks unguarded on about the same calendar date, regardless of chick age. However, chicks from long incubation nests averaged 8 days younger when they were left unguarded than chicks from control or short-incubation nests. In females, there was no effect of nest failure on prolactin levels. In males, prolactin levels were slightly lower after nest failure than in males tending nests. Testosterone was significantly higher in males after nest failure than in males still tending nests. Prolactin is elevated in Adélie penguins as part of the program of cyclical hormonal changes that accompany the lengthy reproductive season and is relatively independent of tactile input. Sustained prolactin secretion is probably required for the maintenance of parental behavior in offshore feeding species that must be absent from the nest for many days at a time.     

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.     

Feeding territoriality and the reproductive consequences in brown skuas<Emphasis Type="Italic"> Catharacta antarctica lonnbergi</Emphasis>

In the maritime Antarctic, brown skuas (Catharacta antarctica lonnbergi) show two foraging strategies: some pairs occupy feeding territories in penguin colonies, while others can only feed in unoccupied areas of a penguin colony without defending a feeding territory. One-third of the studied breeding skua population in the South Shetlands occupied territories of varying size (48 to >3,000 penguin nests) and monopolised 93% of all penguin nests in sub-colonies. Skuas without feeding territories foraged in only 7% of penguin sub-colonies and in part of the main colony. Females owning feeding territories were larger in body size than females without feeding territories; no differences in size were found in males. Territory holders permanently controlled their resources but defence power diminished towards the end of the reproductive season. Territory ownership guaranteed sufficient food supply and led to a 5.5 days earlier egg-laying and chick-hatching. Short distances between nest and foraging site allowed territorial pairs a higher nest-attendance rate such that their chicks survived better (71%) than chicks from skua pairs without feeding territories (45%). Due to lower hatching success in territorial pairs, no difference in breeding success of pairs with and without feeding territories was found in 3 years. We conclude that skuas owning feeding territories in penguin colonies benefit from the predictable and stable food resource by an earlier termination of the annual breeding cycle and higher offspring survivorship.Research licence: Umweltbundesamt Bonn 13.4-94003-1/5-7.     

Blackcap Sylvia atricapilla nestlings do not produce begging calls until they are able to escape from predators

Nest predation is a major source of reproductive failure in birds and thus it can exert selection on both parental and offspring strategies. Begging calls are known to be a powerful component of parent–offspring communication but these calls can also increase predation risk. Here we demonstrate a sophisticated strategy for the development of begging vocalization in a species under high nest predation. Blackcap Sylvia atricapilla nestlings spend most of their nesting period silent, and develop begging calls just before they are able to fledge. The onset of begging vocalization matches the onset of endothermy, which enables Blackcap chicks to leave the nest. We demonstrate experimentally that begging calls function as a signal of the increased needs of homeothermic nestlings. Playback of begging calls conducted in nests with silent nestlings resulted in a significant increase in feeding rates and a decrease in brooding. Development of begging calls only at the age of endothermy allows species under high nest predation to keep the risky period of begging vocalizations and frequent feeding to a minimum. This strategy may constitute an evolutionary solution to high predation pressure in some open nesting passerines. This is the first study to demonstrate the existence of silent begging in a passerine.     

Acoustic Communication in a Black‐Headed Gull Colony: How Do Chicks Identify Their Parents?

Chicks perform conspicuous begging behaviour in response to the arrival of a parent. In seabirds colonies, as nests are close to each other, chicks are permanently surrounded by sound and visual stimuli produced by adult conspecifics approaching their nests. However, in spite of these conditions, black-headed gull chicks begin to vocalize as their parent approaches even before they can see it. In this paper, we report field experiments testing sound-based discrimination of parents by black-headed gull chicks. Focusing on the 'long call', i.e. the signal emitted by parents when coming back to the nest, we investigate here the acoustic parameters used for this recognition process. By playback experiments using modified 'long calls', we demonstrated that signals without amplitude modulation still elicit responses in chicks. In contrast, frequency modulation appears essential. In the frequency domain, experiments revealed that frequency analysis is precise. Chicks did not react when the frequency spectrum of parental call was shifted 20 Hz down or up. The totality of harmonics is not necessary: chicks require only two harmonics to discriminate between parents. Signal redundancy is of great significance since a minimum of four successive syllables in parental 'long call' are required to elicit reaction in the chick.     

An observation of between-mates feeding behaviour in chick-guarding chinstrap penguins

Courtship feeding has not been reported for chinstrap penguins (Pygoscelis antarctica). However, we observed a male chinstrap penguin fed to his mate during the guard stage of chick. After being fed, the female regurgitated the food to her chick. Our observation suggests that the chicks’ behaviour of begging for food may be retained in adults, but it is usually restrained.     

Parent-offspring interactions in food provisioning of Manx shearwaters: implications for nestling obesity

Procellariiform seabirds such as the Manx shearwater Puffinus puffinus, rear only one chick at a time but may breed many times in their lives; parents should thus limit food delivery to the chick in keeping with the balance between current and future reproductive output. Yet procellariiform chicks accumulate large quantities of lipid, which may provide a buffer against pronounced and unpredictable variation in food provisioning, resulting in part from an inability of parents to regulate food supply to the nest. We switched chicks between nests to examine the roles of parents and offspring in controlling food delivery. The serial autocorrelation in age-specific body masses for unmanipulated chicks decreased from 0.61 (P< 0.01) to 0.35 (NS) over a period of 15 days and remained nonsignificant thereafter. By contrast, the serial autocorrelation for switched chicks increased from 0.64 (P< 0.01) to 0.83 (P< 0.001) and the serial cross-correlation rose from 0.23 (NS) to 0.50 (P< 0.05). These results supported both chick determination and parental determination models of food provisioning, indicating that chicks conveyed information about their nutritional status, which parents acted upon by adjusting their rate of food delivery. We discuss these results in relation to the optimization of nestling lipid reserves and parental foraging effort. We suggest that information conveyed by the chick's begging intensity serves to reduce the provisioning rate to well-fed chicks, but parents cannot or do not increase food provisioning to poorly fed chicks. Such adjustment of food provisioning does not refute the hypothesis that nestling obesity provides a buffer against highly variable food delivery. Copyright 1999 The Association for the Study of Animal Behaviour.     

Provisioning behaviour at the nest in single-parent versus biparental nests and male versus female parents in the common redstart (<Emphasis Type="Italic">Phoenicurus phoenicurus</Emphasis>)

We analysed video-sequences of undisturbed parental provisioning behaviour on 12 nests of common redstart (Phoenicurus phoenicurus). In 4 of the 12 nests, chicks were fed by a single parent only. We compared provisioning rate of chicks, time spent on the nest and food allocation rules between nests with uniparental and biparental care and between male and female parents in biparental nests. In nests with a single parent, the frequency of feeding visits per parent was higher than in biparental nests. As a result, the rate of food provisioning of chicks was similar in uniparental and biparental nests. The food allocation rules did not differ between uniparental and biparental nests. In biparental nests, male and female provisioning behaviour was similar though with two exceptions: males had a strong preference for feeding chicks in front positions in the nest and females spent a longer time on the nest after feeding. We conclude that single common redstart parents are able to compensate fully for the absence of the other parent through increased provisioning efforts, and that in biparental nests, males and females contribute equally to the provisioning of the young.     

Adult aggression during the post-guard phase in the chinstrap penguin Pygoscelis antarctica

We report an observational study of aggressive behaviour by adults during the post-guard phase in the chinstrap penguin. The study was carried out in one subcolony where all the individuals (breeders, chicks and failed breeders) were banded, as well as in nine other subcolonies where individuals were not identifiable. Breeding adults were more aggressive towards unrelated chicks when they were feeding their own chicks than in other contexts. Chicks were also attacked at high rates by adults that did not belong to their own subcolonies. The evidence suggests that some aggressions to chicks by adults may function to avoid interference from unrelated chicks during food transfers, but our data also show that aggressive interactions are common in contexts other than chick feeding. However, aggression by adults did not show any clear spatial directionality or purpose of shepherding the chicks, and these tended not to move in any particular direction when attacked.     

Regulation of parental investment in the Antarctic petrel Thalassoica antarctica: an exchange experiment

An experiment was conducted on the Antarctic petrel to test whether the parents were able to respond to changes in food demand of their offspring. Two experimental groups were formed by replacing eight 20-day-old chicks with 10-day-old chicks, and vice versa. The growth rate of chicks in the experimental groups was compared with that in two control groups with chicks of known age. The growth rate of 10-day-old chicks in the nests of parents which initially had 20-day-old chicks did not differ significantly from that in their respective control groups. This indicates that those parents were able to raise a new young nestling, despite having already raised another chick from hatching to 20 days. However, the 20-day-old chicks placed in nests with 10-day-old chicks had a significantly lower growth rate than their control group. Feeding rate per day and nest did not differ significantly among any of the groups. This suggests that the observed difference in growth rate between 20-day-old chicks is related to a lower amount of food delivered per visit to experimental chicks. Thus, in the Antarctic petrel, the feeding rate apparently is not regulated by the status of the chick, but by the parents' ability to gather food or willingness to provide food for the chicks.Publication no. 137 from the Norwegian Antarctic Research Expedition (NARE) 1989/90     

Development of chicks and predispersal behaviour of young in the Eagle Owl Bubo bubo

Little quantitative information on the development and behaviour of chicks and young is available for many species, despite the crucial importance of such data and the sensitivity of this stage in a bird's life. For Eagle Owls Bubo bubo , despite the large amount of scientific literature on this species, much basic information is lacking. This study provides a photographic and morphometric guide for age estimation of nestlings and fledglings, as well as data on the call behaviour of young, and patterns of movements during the post-fledging dependence period. The most remarkable event in chick development is the rapid increase in mass, and size gain, during the first 30 and 40–45 days, respectively. Because after this time morphometric differences become less evident, young-feather development is more useful for ageing. Patterns of chick call behaviour showed that the time spent calling increased with age and, from 110 days of age, chick vocalizations were usually uniformly distributed through the whole night and most synchronized at sunset and sunrise (the maximum recorded number of vocalizations per chick and per night was 1106 calls). During the post-fledging dependence period, radiotagged Owls moved widely, up to 1500 m from the nest after the age of 80–90 days. During such movements, the mean distance among siblings increased with age, from 168 m on average for juveniles less than 100 days old, to 489 m for those older than 100 days. Definitive dispersal started when young were about 150–160 days old. Information on chick call behaviour and movements is crucial for unbiased census and nest checking, as well as for the definition of young post-fledging areas. Knowledge of the latter is very important in terms of conservation and management (especially for those species that move largely around their nest before dispersal) owing to the high mortality that can occur during this period.     

Does larder-hoarding affect the condition of chicks in urban kestrels?

Hoarding by urban kestrels and its subsequent exploitation significantly increases the frequency and regularity of feeding during the early stage of the nestling period. Twelve kestrels’ nests situated in various part of the city were chosen to test whether hoarding could affect the chicks’ condition. Four nests (with 22 nestlings) were provided with food — one dead mouse per chick per day (F group) and 8 (43 nestlings) were used as a control group (NF). Nestlings were weighed daily, and the length of their tarsus was also measured when the oldest chick in the nest was 11 days old. 50% of the NF nests experienced a decrease in the number of nestlings but no nestlings were lost in any of the F nests. However, no significant differences were found between the groups either in body mass or tarsus length. A possible reason was the behaviour of the adults which could be interpreted as the regulation of the frequency of prey delivery depending on its presence or absence in the nest.     

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.     

Chick loss in the Falkland Skua Catharacta skua antarctica

Data on reproductive success of 110 Falkland Skua Catharacta skua antarctica pairs were gathered during the austral summers of 1988–1989 and 1990–1991 on New Island, Falkland Islands. Adults laid two eggs 2–3 days apart and began incubation with the first egg. For the years combined, 1.39 chicks per nest hatched and 0.84 chicks per nest fledged (fledging was defined as surviving to 16 days of age). Brood reduction was common; 43% of the two-chick broods were reduced to one, and mortality was concentrated on the younger chick. Although asynchronous hatching and differential death are consistent with Lack's brood reduction hypothesis, application of O'Connor's quantitative criterion revealed that sibling competition may not be responsible for the observed chick mortality. Furthermore, because no aggressive interactions between chicks were observed or detected indirectly, siblicide may be absent in this population. Instead, predation modified by a variety of factors may have led to the greater mortality of the second-hatched chick.     

Begging coordination between siblings in Black-headed Gulls

Communication behaviours are now considered from a signallers–receivers network perspective. This concept seems well suited to the study of interactions between parents and offspring in birds, so far mainly treated as a dyadic signalling system involving the brood or a single chick as a signaller and the parent as a receiver. Family conflicts over resource allocation drive parent–offspring and sib–sib communication. In the Black-headed Gull Larus ridibundus, parents respond to the whole-brood begging intensity and siblings often synchronize their begging signalling thus limiting individual effort. By monitoring five nests of two-chick broods during the whole rearing period in the nest, we show how an intra-brood simultaneity of begging emerges from successive phases of solitary begging of junior and senior nestlings. Although this result remains preliminary due to the sample size, it underlines a dynamical aspect of chicks’ behaviour. Because they always favour coordinated begging and because they elevate their response threshold across the rearing period, parents may play a major role in the plasticity of begging behaviour.     

OBSERVATIONS ON THE BREEDING OF THE PINK-BACKED PELICAN PELECANUS RUFESCENS

This paper summarises observations on the breeding behaviour of the Pink-backed Pelican Pelecanus rufescens at Rakewa, Nyanza Province, Kenya, where the species has bred for at least 200 years. Observations covered most of one breeding season, November 1962 to April 1963. Of at least 250 nests, 35 were closely observed. The community consisted of about 815 pelicans of which about 540 were adults. The death rate is estimated at 13% per annum and the mean life-span at about seven and a half years. The breeding site, in trees above a small swamp, is 15 miles from the favoured feeding ground. The colony is protected by local Luo people. The pelicans feed and roost mainly at the Miriu Delta, 15 miles away, travelling between the two places so high up as to be unseen. They fish in the early morning, visiting the colony to feed young mainly between 09.00 and 13.00 hrs. Once the young are large both parents roost away from the colony at the Delta. The breeding season takes place from August, towards the end of the rains, to March, at the end of the dry season. The birds breed in synchronised groups, the breeding cycle for any group occupying five months. Nuptial display is performed on the nest trees, by single pairs or small groups. Two main displays are described, “pointing” and “bill-clapping”. Mating occurs on the nest, with little preliminary display. Nests are slight stick structures, repaired from year to year, and used by other pelicans if abandoned. The clutch is normally two eggs, occasionally three. Both sexes incubate, with infrequent change-overs, for 33–35 days. The chick is first brick-red, becoming covered with white down. Feathers break through at about 12 days and have covered much of the body by 30 days. At 40 days chicks can recognise their own parent. They fly at 70–75 days. Parents feed chicks by regurgitation, sometimes into the nest. They brood them closely at first, but after 10–12 days leave them much alone. Large chicks thrust the head far into the parental gullet, and injuries result from such feeding struggles. Feeding usually occurs before mid-day, each parent normally delivering two feeds with a rest between. Curious convulsive movements of the young are probably begging displays. Forty-two young hatched in 35 nests, an average of 0.6 chicks/egg laid. The heaviest mortality among young occurred between 10–30 days when 31% of all chicks died. Young which flew were produced at the rate of 0.47/egg hatched, 0.28/egg laid, and 0.57/pair.     

Maternal testosterone influences a begging component that makes fathers work harder in chick provisioning

In species with biparental care, parents disagree evolutionarily over the amount of care that each of them is willing to provide to offspring. It has recently been hypothesised that females may try to manipulate their mates by modifying offspring begging behaviour through yolk hormone deposition, shifting the division of labour in their own favour. To test this hypothesis we first investigated how yellow-legged gull (Larus michaellis) parents feed offspring in relation to each component of complex begging behaviour and if feeding behaviour varies between sexes. Then we investigated the effect of yolk testosterone on chicks' begging by experimentally increasing yolk testosterone levels. Our results revealed that yolk testosterone has a component-specific effect on chicks' begging, specifically increasing the number of chatter calls. Parental feeding effort was influenced by the number of chatter calls emitted by chicks, but most importantly, the influence was stronger in male than in female parents. Moreover, chick body mass increased with the number of paternal feeds. In conclusion, these results show that female gulls may use yolk testosterone deposition to exploit their partners as predicted by the ‘Manipulating Androgen Hypothesis (MAH)’.