The response of willow grouse chicks to auditory stimuli 1. Preference for hen grouse calls

Willow grouse (Lagopus lagopus lagopus) chicks newly hatched from mechanically incubated eggs were exposed to different combinations of sounds in a heated runway. They approached a loudspeaker playing grouse hen calls, which they had never heard, in preference to one playing the noise of the hatching machine from which they had just been removed. Similarly they preferred the hen call to another novel sound. When the three sounds were played singly instead of in pairs, only the hen call was approached.     

Development of responsiveness to the individual maternal “clucking” by domestic chicks (Gallus gallus domesticus)

Sixty-nine chicks, brooded by hens, were given the choice of approaching the clucking sound of the familiar hen or another clucking source. The effects of age and prenatal experience of clucking on the responsiveness to their hen's clucking were also investigated. Eighty-three percent of all chicks tested chose the familiar hen clucking after a certain age. By 4 days of age, 29% of artificially incubated chicks approached the clucking of a familiar hen. Seventy-four percent of naturally incubated chicks and 89% of chicks artificially incubated until pipping and then naturally incubated, showed this discrimination. This corresponds well to the natural situation, in which the hen remains in the nest with the chicks for up to 3 days after hatching. The results also indicate that late prenatal experience of the maternal call is of importance for the later responsiveness to this call.     

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.     

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.     

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.     

Adult chicken alarm calls enhance tonic immobility in chicks

An experimenter induced tonic immobility (TI) in parentally naive chicks (G. gallus domesticus). The chicks remained in TI longer when they were exposed to a conspecific adult fear squawk alarm call than when exposed to an equally novel attraction call or white noise. In a second experiment, both aerial-predator and ground-predator alarm calls enhanced TI similarly to the fear squawk call which is elicited by capture. These results support the hypotheses that TI is an antipredator defense mechanism and that alarm calls evolved through kin-selection.     

On the responses of laughing gull chicks (Larus atricilla) to the calls of adults II. Age changes and responses to different types of call

Three age-groups of laughing gull chicks were tested with playback of recordings of calls of their parents and calls of neighbouring adults. The chicks in each group showed response to the playback calls, and recognition of the voices of their parents. The chicks in the early group (1 to 3 days post-hatching) showed the least evidence of parent recognition: the chicks of the late group (12 to 28 days post-hatching) showed the lowest levels of response in general.Evidence of relationships between variation in the behaviour in the tests and variation in the test recordings was found from correlation analysis, the type of call initiating positive response, and analysis of temporal association. This evidence, and the results of tests which allowed responses to the parents' crooning and responses to the parents' ke-hah calls to be compared for chicks in the late group, indicate that whereas the filial approach behaviour of very young chicks is stimulated by crooning but suppressed by ke-hah and long-calls, this behaviour in older chicks requires the occurrence of ke-hah calls or long-calls for its elicitation. However the ke-hah calls and long-calls of adults other than the parents continue to have negative effects on the behaviour of older chicks.It is argued that this progression in discrimination of the voices of the parents and the changes in responsiveness to the different types of adult calls are consistent with functional requirements, both from the point of view of changes in a growing chick's social situation and from the point of view of the characteristics of the calls. The changes are also discussed from a developmental point of view.     

Interactions between adult robins and chicks belonging to other pairs

Adult robins are able to discriminate between chicks hatched in their own and in other territories. The latter are much less likely to be fed and are more likely to be attacked than the adult's own young. As chicks grow older they become less likely to beg at adults other than their parents. Adults rarely feed chicks from other territories unless they have fledged young of their own. Possible causes of these different responses are discussed and it is emphasized that these need not involve individual recognition. Males appear to be more likely than females to feed chicks hatching in other territories. This could be the result of the females learning about their chicks while brooding them. Males may benefit more, on average, from feeding other pairs' chicks than do females. There has been a previous report that robins are unable to make the discriminations reported in this paper and possible reasons for this discrepancy are discussed. It is conceivable that it reflects a difference between populations.     

Responses of ring-necked pheasant chicks (Phasianus colchicus) to conspecific calls

The locomotor responses of pheasant chicks to conspecific calls and the vocalizations of the chicks relating to their locomotor responses were measured in a test apparatus permitting approach or retreat movements. The brood-gathering call and the content call elicited strong approach from chicks. The alarm call and the squeak call inhibited locomotion in chicks. The hiss call, brood-caution call, fright call and flock call do not greatly affect the locomotion of subjects. The flock call appears to be primarily a care-soliciting call. Conspecific calls appear to be important to the survival of young pheasant chicks in the wild.     

An investigation into egg-acceptance by azure-winged magpies and host-recognition by great spotted cuckoo chicks

Tests using magpie's eggs (which are very similar to those of the great spotted cuckoo) on azure-winged magpies (Cyanopica cyanea) showed that the latter accept strange eggs, rejecting only 25% of them. We therefore suggest that they are an accepter species, and as such a substitute host species for the great spotted cuckoo (Clamator glandarius). We suggest a theory for the transition from ‘Accepter Species’ to ‘Non-mimetic-egg Rejecter Species’ by hosts of specialist brood parasites. We have obtained evidence of host recognition by great spotted cuckoo chicks of typical or atypical host species characteristics. Basing our theory on the calls of parasitic chicks reared in the nests of the azure-winged magpies and magpies (Pica pica), we suggest that the foregoing is the mating mechanism necessary for the parasitization of a new host species.     

Vocalizations of chicks and juveniles and the development of adult calls in the Aldabra White-throated rail Dryolimnas cuvieri aldabranus (Aves: Rallidae)

Calls were recorded from eggs, chicks and juveniles of the Aldabra White-throated rail Dryolimnas cuvieri aldabranus. A sonagraphic analysis was made of these calls, their behavioural contexts described and probable functions suggested. Three calls, the twitter, tiuu and contented peep were produced by chicks in the egg. All three calls may be derived from the basic peep first recorded some 36 hours before hatching. Shortly after hatching two more calls, the distress call and the alarm call, can be elicited from the rail chicks. Contented peeps and twitters are restricted to the repertoire of young rails. Song was first heard from wild rail chicks at the age of ten days and may develop from the contented peep. Three adult calls, the 'mp yeah, 'mptiuu and 'mpclick appear at the age of three months and six months later the adult vocal repertoire is completed by the appearance of 'mps, toks, purrs and nest-defence squeals. With the exception of the 'mp, all adult calls can be derived from the vocalizations of chicks and juveniles.     

Oxygen consumption rates of adults and chicks during brooding in king quail (Coturnix chinensis)

Oxygen consumption rates were measured in chicks (0–7 days of age), and in non-brooding and brooding adults. Brooded chicks maintained a constant oxygen consumption rate at a chamber ambient temperature of 10–35°C (0–5 days of age: 2.95ml O₂·g^-1·h^-1 and 6–17 days of age: 5.80 ml O₂·g^-1·h^-1) while unbrooded chicks increased oxygen consumption rate at ambient temperature below 30°C to double the brooded oxygen consumption rate at 25 and 15°C for chicks < 5 days of age and>5 days of age, respectively. The massspecific oxygen consumption rate of breeding male and females (non-brooding) were significantly elevated within the thermoneutral zone thermal neutral zone (28–35°C) in comparison to non-breeding adults. Below the thermal neutral zone, oxygen consumption rate was not significantly different. The elevation in oxygen consumption rate of breeding quail was not correlated with the presence of broodpatches, which developed only in females, but is a seasonal adjustment in metabolism. Male and females that actively brooded one to five chicks had significantly higher oxygen consumption rate than non-brooding quail at ambient temperature below 30°C. Brooding oxygen consumption rate was constant during day and night, indicating a temporary suppression of the circadian rhythm of metabolism. Brooding oxygen consumption rate increased significantly with brood number, but neither adult body mass nor adult sex were significant factors in the relationship between brooding oxygen consumption rate and ambient temperature. The proportion of daylight hours that chicks were brooded by parents was negatively correlated with ambient temperature. After chicks were 5 days old brooding time was reduced but brooding oxygen consumption rate was unchanged. Heat from the brooding parent appeared to originate mainly from the apteria under the wings and legs rather than the broodpatch. The parental heat contribution to chick temperature regulation below the chicks' thermal neutral zone is achieved by increasing parental thermal conductance by a feedback control similar to that suggested for the control of egg temperature via the brood-patch. It is concluded that the brooding period is an energetic burden to parent quail, and the magnitude of the cost increases directly with brood number and inversely with ambient temperature during this period. The oxygen consumption rate of brooding parents was 5.80–6.90 ml O₂·g^-1·h^-1 (ambient temperature 10–15°C) at night and up to 5.10 ml O₂·g^-1·h^-1 (ambient temperature 18°C) during the day, which are 100 and 40% higher than non-brooding birds, respectively.Abbreviations bm body mass - SMR standard metabolic rate - T _a ambient temperature - T _b body temperature - I/O₂ oxygen consumption rate - C _wet wet thermal conductance - TNZ thermal neutral zone - ANOVA analysis of variance - ANCOVA analysis of covariance     

Acoustic systems are adapted to breeding ecologies: individual recognition in nesting penguins

In all species of penguins studied to date, the display call, or parental call, has been demonstrated experimentally to facilitate identification between mates and between chicks and parents. We investigated parent-chick recognition in two nesting species, the Adélie penguin, Pygoscelis adeliae, and the gentoo penguin, P. papua. Through playback experiments, we tested the capacity of chicks to recognize the parental call at varying levels of background noise. By using modified calls, we found that chicks of neither species used temporal characteristics of the parental call (variations in frequency or amplitude with time) for individual recognition, but that both species used a simpler parameter, the pitch of the call. This finding contrasts with the more sophisticated use of acoustic cues by chicks of two non-nesting species, identified in earlier work. These differences in auditory processing of parental calls may have evolved because of different ecological constraints, particularly whether recognition of a nest site supports mutual identification of parent and offspring. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Chick Begging Calls Reflect Degree of Hunger in Three Auk Species (Charadriiformes: Alcidae)

Begging behaviour is an important element in the parent-offspring conflict; it has been studied in many avian species. However, the majority of the studies have been entirely based on the call counts, and they agreed that vocal activity was a good indicator of chick’s nutritional need and/or condition. Fewer researches were dedicated to the temporal-frequency variables of the begging calls themselves and they showed contrary results. Here begging behaviour in three burrow nested, uniparous species of auks (Alcidae) was studied. These objects provide an opportunity to study the signalling value of begging calls in the absence of important confounding factors such as nestling competition and predation pressure. I recorded calls of individual chicks in two conditions: during natural feeding and after experimental four-hour food deprivation. I found that almost all measured acoustic variables contain information about the chick’s state in all studied species. The hungry chicks produced calls higher in fundamental frequency and power variables and at higher calling rate compared to naturally feeding chicks. The effect of food deprivation on most acoustic variables exceeded both the effects of individuality and species. In all studied species, the frequency variables were stronger affected by hunger than the calling rate and call durations. I suppose that such strong change of acoustic variables after food deprivation can be explained by absence of vocal individual identification in these birds. As parents do not need to check individuality of the chick in the burrow, which they find visually during the day time, the chicks could use all of the acoustic variables to communicate about their nutritional needs.     

Can acaricide‐impregnated leg bands fitted to female red grouse reduce sheep tick parasitization of chicks and increase chick survival?

In parts of northern England, North Wales and the Scottish Highlands, increasing numbers of sheep ticks Ixodes ricinus (Ixodida: Ixodidae), and the louping ill virus they can carry, are considered to be important factors that reduce red grouse Lagopus lagopus scotica productivity. The present study tested this hypothesis by fitting adult female grouse with leg bands impregnated with the acaricide cypermethrin to experimentally control ticks on their chicks on two managed grouse moors in northeast Scotland. The chicks of females fitted with acaricide leg bands showed reduced tick infestations and improved survival in one of the two study years, relative to chicks of control females. Acaricide leg bands constitute a potential management technique that may be adopted by grouse moor managers in circumstances of high tick infestations on grouse chicks.     

Ultradian activity rhythms in large groups of newly hatched chicks (Gallus gallus domesticus)

A clutch of young chicks housed with a mother hen exhibit ultradian (within day) rhythms of activity corresponding to the brooding cycle of the hen. In the present study clear evidence was found of ultradian activity rhythms in newly hatched domestic chicks housed in groups larger than natural clutch size without a mother hen or any other obvious external time-keeper. No consistent synchrony was found between groups housed in different pens within the same room. The ultradian rhythms disappeared with time and little evidence of group rhythmicity remained by the third night. This disappearance over time suggests that the presence of a mother hen may be pivotal for the long-term maintenance of these rhythms. The ultradian rhythm of the chicks may also play an important role in the initiation of brooding cycles during the behavioural transition of the mother hen from incubation to brooding. Computer simulations of individual activity rhythms were found to reproduce the observations made on a group basis. This was achievable even when individual chick rhythms were modelled as independent of each other, thus no assumptions of social facilitation are necessary to obtain ultradian activity rhythms on a group level.     

Imperfect mimicry of host begging calls by a brood parasitic cuckoo: a cue for nestling rejection by hosts?

Coevolutionary interactions between avian brood parasites and their hosts often lead to the evolution of discrimination and rejection of parasite eggs or chicks by hosts based on visual cues, and the evolution of visual mimicry of host eggs or chicks by brood parasites. Hosts may also base rejection of brood parasite nestlings on vocal cues, which would in turn select for mimicry of host begging calls in brood parasite chicks. In cuckoos that exploit multiple hosts with different begging calls, call structure may be plastic, allowing nestlings to modify their calls to match those of their various hosts, or fixed, in which case we would predict either imperfect mimicry or divergence of the species into host-specific lineages. In our study of the little bronze-cuckoo (LBC) Chalcites minutillus and its primary host, the large-billed gerygone Gerygone magnirostris, we tested whether: (1) hosts use nestling vocalizations as a cue to discriminate cuckoo chicks; (2) cuckoo nestlings mimic the host begging calls throughout the nestling period; and (3) the cuckoo begging calls are plastic, thereby facilitating mimicry of the calls of different hosts. We found that the begging calls of LBCs are most similar to their gerygone hosts shortly after hatching (when rejection by hosts typically occurs) but become less similar as cuckoo chicks get older. Begging call structure may be used as a cue for rejection by hosts, and these results are consistent with gerygone defenses selecting for age-specific vocal mimicry in cuckoo chicks. We found no evidence that LBC begging calls were plastic.     

Effects of removal of chicks from hens on concentrations of prolactin, luteinizing hormone and oestradiol in plasma of brooding Gifujidori hens.

Gifujidori hens were allowed to repeat a breeding cycle in one season. In the first breeding cycle the duration of the brooding (raising chicks) stage was limited to 3 weeks, whereas in the second breeding cycle it was limited to 1 week by removing all chicks from mother hens. In the first breeding cycle, plasma prolactin (PRL) was high during the incubation period, but rapidly decreased on the day of hatching and reached minimum values about 1 week after hatching. In contrast, plasma luteinizing hormone (LH) concentrations were low during the incubation period, but after hatching they gradually increased and reached peak values immediately after removal of chicks. Concentrations of oestradiol in plasma were low in the incubation and brooding stages but increased significantly immediately after removal of chicks. In the second breeding cycle, changes in PRL and LH concentrations were similar to those observed in the first breeding cycle except that even greater increases in plasma LH and oestradiol concentrations were observed one week after hatching when the chicks were removed. These results suggest that coexistence of newly hatched chicks may suppress LH secretion from the pituitary of the hen in the natural breeding cycle.     

Maternal vocalizations of mallard ducks (Anas platyrhynchos)

While incubating, brooding and calling their young out of the nest, female mallard ducks (Anas platyrhynchos) utter a species-typical maternal vocalization that their young find highly attractive. To determine the characteristic acoustic features of these calls, we recorded the vocalizations of seven hens in the field. The pre-exodus and nest exodus calls of these hens were similar with respect to frequency modulation, presence of a low-frequency impulsive sound, note duration, and repetition rate. The exodus call differs from the pre-exodus call in having more notes per burst and more harmonics, with a corresponding upward shift in dominant frequency. Repetition rate and frequency modulation may be the critical acoustic features of the auditory basis of species identification in mallard ducklings.     

Brooding chicks with or without supplemental heat and light

The need for supplemental heat in brooding chicks in a hot humid tropical area where prevalent mean environmental temperature and relative humidity are 33°C and 60%, respectively, was tested in an experiment using 320 broiler chickens.The chicks were divided into four quadruplicate lots of 20 birds each and were brooded with or without supplemental heat and light for periods varying between zero and 28 days. Each treatment group of 80 birds was subjected to one of four heat and light treatments, namely: supplementation with heat and light for the 28-day brooding period (control), supplementation with heat and light for the first 14-days only, supplementation for the last 14-days only and non-supplementation with heat and light for the entire 28-day period. Final body weight, body weight gain, feed consumption, feed conversion efficiency and mortality were not significantly (P>0.05) affected by the treatments. However, chicks brooded with supplemental heat and light for 28 days consumed significantly more water than all others. Chicks brooded without supplemental heat and light for 28 days had significantly (P<0.05) higher haemoglobin and hematocrit values than all others.