Cooperative breeding in purple gallinules: the role of helpers in feeding chicks

Summary The participation of breeders and helpers in the feeding of 21 broods of chicks was studied in a population of cooperatively breeding purple gallinules (Porphyrula martinica). In the breeding group, all birds over the age of 2 months fed chicks. Female breeders fed chicks at the highest rate, followed by male breeders and adult helpers, old juvenile helpers, and young juvenile helpers. The amount of food breeders fed chicks was independent of the number of helpers in the breeding group. However, breeders made fewer feeding visits when they had helpers. Male and female breeders spent similar amounts of time feeding chicks. Helpers spent significantly less time feeding chicks than did breeders. As helpers grew older they fed chicks at a faster rate, made more feeding visits and spent more time feeding chicks. Analysis of variance was used to determine which variables explained the variation in the brood feeding rate (amount of food delivered to an entire brood during one observation). Age of chicks had a significant nonlinear effect, and size of brood and number of helpers had significant linear effects on the brood feeding rate. Chicks in groups with helpers received more food and were accompanied for longer periods of time than chicks in groups without helpers; either or both of these factors may have led to increased chick survival.     

Cooperative breeding by the Galápagos mockingbird,Nesomimus parvulus

Summary The costs and benefits of helping behavior were analyzed for 36 pairs of the Galápagos mockingbird, Nesomimus parvulus, and their associates. Helping at the nest is usually done by sons or males suspected to be offspring of the breeders. Costs and benefits to breeders were assessed by comparison of pairs with and without helpers, and costs and benefits to helpers were assessed by comparison of birds which help and those which establish themselves as novice breeders.Helping behavior benefits breeders by increasing fledging success and by reducing the adult energy load in territory defense and feeding of nestlings. Breeders assisted by helpers may also benefit by decreased nest predation. Helpers enhance their inclusive fitness by helping, and gain directly by increasing their chances of securing a territory. Helpers do not appear to gain any fitness advantage from the experience of assisting, nor do they increase their survivorship by remaining on natal territory.Ecological and demographic features such as saturated territories and low territory turn-over rates due to high adult survival may be primarily responsible for the evolution of the helping behavior, with kinselection reinforcing it. Associated features of this system are a male-biased population sex ratio, a greater energetic benefit to breeding males than to breeding females in having helpers, earlier dispersal and breeding by females than by males, and much more frequent helping by males than by females. These are interpreted as consequences of brothersister aggression that indirectly minimizes the chances of inbreeding.     

Correlates of group size in a cooperatively breeding cichlid fish (Neolamprologus pulcher)

Neolamprologus pulcher, a cooperatively breeding cichlid fish from Lake Tanganyika, lives in permanent social groups comprising one breeding pair and helpers of both sexes. Variation in group size (1-14 helpers) provides an opportunity to investigate factors that affect how many helpers remain in a group and in turn how group size affects reproductive success. This field study showed that larger groups live in larger territories with more shelter. Group size was more strongly correlated with territory quality than with breeder size. Experimental enhancement of territory quality did not affect group size but group size decreased when territory quality was reduced. Breeders living in a large group benefit because such individuals feed more often and have lower workloads and greater reproductive success. Helpers in larger groups also fed more frequently but did not have lower workloads. This is one of the first experimental studies to examine the factors influencing group size in cooperative breeders.     

Social and mating system of cooperatively breeding laughing kookaburras (Dacelo novaeguineae)

DNA fingerprinting was combined with field observations over four breeding seasons to investigate the social structure and mating system of the laughing kookaburra (Dacelo novaeguineae). Groups comprised a socially dominant pair and up to six helpers of either sex. Helpers were always recruited from young hatched in the group. Territorial inheritance, which is a feature of other cooperative breeders and an oft-cited benefit of philopatry, did not occur. Helpers only attained dominant status in an established group by dispersing into a vacant dominant position in that group. However, helpers could also form new groups by excising a new territory, often through a ”budding” process. The mating system was overwhelmingly monogamous. There were no cases of extra-group parentage in a sample of 140 nestlings; within groups of three or more birds, dominance predicted parentage almost perfectly (99.2% of 129 nestlings), irrespective of whether helpers in the group were related to one or both dominant birds. This is contrary to predictions from models of reproductive skew, possibly because they currently fail to incorporate the willingness of females to share reproduction among males. Received: 15 May 1999 / Received in revised form: 2 November 1999 / Accepted: 6 November 1999     

The role of helpers in feeding chicks in cooperatively breeding green (red-billed) woodhoopoes

Summary Observations were made of ten green (red-billed) woodhoopoe Phoeniculus purpureus flocks during the breeding season in order to quantify the relationship between flock size and the amount of food delivered to chicks. The study period was kept short specifically to minimize the effects of environmental stochasticity. Neither woodhoopoe feeding visit rates nor the total amount of food brought to chicks increased with flock size. Although nonbreeders did not increase the net rate of food provisioning to chicks, they reduced parental input in chick rearing, and hence energy expenditure by the breeding pair. However, over an 8-year study period, which includes data for 144 flock years, this did not result in increased breeding frequency or enhanced survival of breeders. There is thus no evidence that helpers' feeding contributions to young per se influence the indirect fitness of helpers.     

Positive correlation between helpers at nest and nestling immune response in a cooperative breeding bird

In cooperatively breeding species, helpers may contribute to the success of the brood by increasing the number of independent offspring, but also, they may affect offspring condition and, hence, their survival and recruitment into the breeding population. This second type of benefits is rarely included in theoretical models or assessed in field studies. Immune response is a good proxy of individual quality and fitness, and there is good evidence that the performance of the immune system of chicks during the nestling phase is related to their chances of survival and future reproduction. However, no study has so far explored whether helpers at the nest might contribute to enhance immune functioning of nestlings in species with a cooperative breeding system. Here we investigate this issue in the azure-winged magpie (Cyanopica cyanus) and found for the first time a positive correlation between the number of helpers at nest and the cell-mediated immune response of nestlings. This effect was not explained by a general improvement of body condition of chicks because it was independent of individual variation in body mass or tarsus length. Our results suggest that helping can have subtle effects on the quality of offspring that may influence their survival and future reproduction.     

Incubation feeding by helpers influences female nest attendance in the green woodhoopoe,<Emphasis Type="Italic"> Phoeniculus purpureus</Emphasis>

In many bird species, the female alone incubates the eggs, but the male provides her with some food during the incubation period. In cooperatively breeding species, helpers might be expected to assist the breeding male in provisioning the female, but their contribution has been generally ignored. Here, I show that in the green woodhoopoe, Phoeniculus purpureus, the breeding male and helpers of both sexes bring food to the incubating female. The helpers did not increase the overall amount of provisioning the female received: groups of all sizes delivered a similar amount of food per hour. Helpers did, however, reduce the workload of the breeding male. Male and female helpers provisioned incubating females equally, as expected in a species where both sexes are likely to derive equal benefits from their helping behaviours. Female nest attendance was positively related to the level of group provisioning, but only in the short term. Thus, the female-nutrition hypothesis, which suggests that incubation feeding can provide an important source of energy to the breeding female, is supported in the green woodhoopoe.Communicated by I. Hartley     

Social system and reproduction of helpers in a cooperatively breeding cichlid fish (Julidochromis ornatus) in Lake Tanganyika: field observations and parentage analyses

It is suggested that some fish of the genus Julidochromis, substrate-brooding Tanganyikan cichlids with biparental care, breed cooperatively with helpers. We studied the social system of J. ornatus in the wild and analysed genetic parentage using microsatellites. Within the studied population three patterns of social system were identified: monogamous pairs (61%, 44 of 72 groups), pairs with helpers (29%, 21), and polygamous harems with helpers (controlled by either a large female or large male owner; 10%, 7). In cooperatively breeding groups, the number of helpers at each nest ranged from 1 to 6 (median 1), and male helpers were more numerous than female helpers. In both sexes, the body size was different among individuals of different social status (harem owner > breeder > helper). Helpers and harem owners of both sexes exhibited brood defence although its frequency was low. The molecular analysis revealed that (1) the helpers were mostly unrelated to dominant breeders, (2) many helpers of both sexes contributed genetically to the next generations, (3) male helpers had high siring success (41% of young in total), and (4) large young unrelated to group members were detected at 30% of observed nests, which may be due to breeder (or helper) replacements and immigration of young. We discuss the implications of these results for understanding the complex social system of this species, especially the low reproductive skew in comparison with other cooperatively breeding cichlids.     

Helpers in colonial cooperatively breeding sociable weavers <Emphasis Type="Italic">Philetairus socius</Emphasis> contribute to buffer the effects of adverse breeding conditions

Some studies on the effects of helpers in cooperatively breeding vertebrates show a positive effect of helper presence on reproductive output whereas others find no effect. One possibility for this discrepancy is that helpers may have a positive effect when breeding conditions are adverse, while their effect might go unnoticed under good conditions. We investigate this hypothesis on sociable weavers Philetairus socius, a colonial cooperatively breeding passerine that inhabits a semi-arid region where breeding conditions vary markedly. We used multivariate mixed models to analyse the effect of helpers on reproduction under contrasting environmental and social conditions while controlling for parental and colony identity. We found that reproductive success in sociable weavers was primarily influenced by nest predation and rainfall. In addition, colony size was negatively associated with hatching and fledging success and number of young fledged per season. Helpers had a less prominent but significant influence on feeding rates and reproductive outcome. In agreement with expectations, the presence of helpers counteracted some of the negative effects of breeding in periods of low rainfall or in large colonies and was also associated with an increased number of young fledged per season. Our results illustrate that the effect of helpers might be detectable mostly under unfavourable conditions, but can contribute to improve reproductive performance in those situations.     

Experimental evidence for helping and hindering by previous offspring in the cooperative-breeding Seychelles warbler Acrocephalus sechellensis

Prebreeding Seychelles warblers (Acrocephalus sechellensis) frequently act as helpers on their natal territory, aiding in territory defence, predator mobbing, nestbuilding, incubation (only females) and feeding dependent young of their parents. In some cases helpers could attain breeding status (e.g. by joint-nesting) in their natal group and become co-breeders. Comparisons of group size and reproductive success on a given quality territory suggest that the presence of alloparents (helpers and cobreeders) significantly affects the reproductive success of their parents. The influence of alloparents on reproductive success was examined by removing alloparents from breeding units and comparing the success of natural-sized and artificially reduced groups. Removal experiments, controlled for territory quality, group size and breeder age, showed that the presence of one alloparent significantly improved the reproductive success of its parents. Analysis strongly suggests that this was entirely due to helping behaviour (i.e. providing care to offspring of their parents), thereby improving the helper's inclusive fitness benefits from staying at home. However, these experiments showed also that the presence of two or more alloparents in medium-quality territories significantly decreased reproductive success, compared with groups with one alloparent. Several lines of evidence suggest that this may have been due to the joint-nesting and reproductive competition that could occur in breeding groups, or simply to resource depression when a large number of previous offspring remained on their natal territory.     

When do helpers help? Food availability and helping in the moorhen,Gallinula chloropus

Summary The importance of food availability as a constraint upon cooperative behaviour was examined by providing supplementary food on some breeding territories of a population of moorhens (Gallinula chloropus). First brood juveniles on territories provided with food were more likely to feed their younger second brood siblings. Supplementary feeding improved the physical condition of juveniles above that of unfed conspecifics, and birds in better condition were more likely to be helpers. Helpers on fed territories also provided more food to siblings than did their counterparts on unfed territories, enabling parent birds to reduce their work rates. Results are discussed in relation to the benefits which may accrue to helpers and the energetic costs of cooperation which may act as proximate constraints upon potential helpers.     

Multiple paternity in the cooperatively breeding fish <Emphasis Type="Italic">Neolamprologus pulcher</Emphasis>

In cooperative breeders, mature males may compete for fertilizations. In this study, we measured the degree of multiple paternity in a natural population of a cooperatively breeding fish. Neolamprologus pulcher (Perciformes: Cichlidae) is a highly social cichlid endemic to Lake Tanganyika. We used highly variable microsatellite loci to survey 12 groups with an average number of 10.6 brood care helpers per group and a total of 43 offspring (mean 3.6 per brood). In 11 of 12 groups, all young were assigned to the dominant female. The dominant male sired all offspring in three groups, part of the offspring in four groups, and in five groups, he had no paternity at all. In total, 44.2% of young were not fathered by the current male territory owner. Multiple paternity was found in 5 of 12 broods (41.7 %), with 8 of 35 young (22.9 %) being sired by males other than the respective territory owners. This is an exceptionally high rate of extra-pair paternity among cooperatively breeding vertebrates. Neither helpers present in these territories during collection nor neighbouring males were unequivocally assigned to have sired these extra-pair young. However, behavioural observations suggest that male helpers may have produced these young before being expelled from the territory in response to this reproductive parasitism. We discuss these results in the light of reproductive skew theory, cooperative breeding in vertebrates and alternative reproductive tactics in fish.     

Sociality and cooperative breeding of red-cockaded woodpeckers,Picoides borealis

Summary Thirty groups of red-cockaded woodpeckers (Picoides borealis) were studied from 1976–1982 to (1) determine the demographic structure of groups, (2) identify the role helpers play in reproductive activities, and (3) investigate the selective pressures promoting sociality and helping behavior. Groups had only 1 mated pair and 0–2 helpers. Approximately half of all groups had helpers and a given group had helpers some years but not others. Helpers, with rare exception, were males 1 or 2 years old and progeny of 1 or both members of the breeding pair. As a result of higher nestling survival, groups with helpers fledged significantly more young per year than unassisted pairs ( , SD=0.97,n=43 vs. , SD=1.01,n=50). Nesting success was also associated with size and quality of nesting period home range, but evidence suggested that the increased number of young fledged resulted directly from the action of helpers. There was a significant positive correlation between reproductive success and experience of breeding females among unassisted pairs but a significant negative correlation among pairs with helpers. In groups with experienced females, helpers were assisting both their mothers and fathers and, therefore, were related to the offspring on the average by 0.50. In groups with inexperienced females, helpers were assisting their fathers and unrelated females and were related to the offspring by 0.25. The red-cockaded woodpecker's unique habit of excavating nest and roost cavities in living pines and the extended period of time required for excavation may be an important ecological constraint that promotes the retention of helpers. Because helpers are related to the offspring they help rear, kin selection and gains in indirect fitness may provide a partial explanation of why red-cockaded woodpecker helpers help. However, the negative correlation between the efficacy of helping behavior and the helpers' relatedness to the offspring they help rear implies that helpers are least effective in producing offspring which would represent greatest gains to indirect fitness. This raises questions about the relative importance of kin selection and indirect fitness in the evolution of helping behavior among red-cockaded woodpeckers.     

Effects of helpers on breeder survival in the red-cockaded woodpecker (Picoides borealis)

Helpers can gain future indirect fitness benefits by increasing the survival of breeders that produce offspring related to the helper. Helping augments group size through the helper's presence and, in some cases, by increasing fledging success. Breeders may then experience enhanced survivorship because of the benefits of living in large groups. Helping may also reduce the workload of the breeder, which in turn may increase the likelihood that the breeder will survive to breed again. We used Cox's proportional hazards model to examine whether breeders' survival in two populations of the red-cockaded woodpecker (Picoides borealis) was enhanced when group size was increased in the presence of (1) the helper itself, or (2) "extra" fledglings (fledglings produced by the breeder because of helping behavior). We found that in the presence of helpers, the risk of a breeder dying declined by 21–42% for males and 0–14% for females. Our results suggest reduced breeder workload as one mechanism to explain reduced breeder mortality in the presence of helpers: breeders spent less time incubating and provisioning nestlings when assisted by helpers. The risk of a breeder dying declined by 16–42% in males and 26–43% in females in the presence of "extra" fledglings. We speculate on possible mechanisms by which fledglings might affect breeder survival. Our results support the hypothesis that helpers gain future indirect benefits by reducing breeder mortality. Electronic Publication     

Helper contributions to reproductive success in the splendid fairy-wren (Malurus splendens)

Summary In Malurus splendens, helpers were present in 65% of 226 group-years with at least one helper female in 37% of group-years. Most females helped for only one year, while many males did so for at least two years. Most were offspring of one or both present breeders, and in 53% of helper-years, helped both parents. For 159 helpers of known age and parentage, the mean coefficient of relatedness to the offspring was 0.47. Novice females with or without helpers produced fewer fledglings per season than females with one year breeding experience and the same level of help. Helpers did not affect production of fledglings per year by females with one year of experience. Females with two or more years experience and at least two helpers produced more fledglings than equivalent birds with one or no helpers. Experience and helpers have little effect on production of fledglings per nest but they lead to more females renesting after a first brood has been raised. Fewer than 20% of novices renest after fledging one brood, while for females with at least two years experience, the percent renesting after success is 40% with no help, 56% with one helper and 69% with 2 or more helpers. Experienced females begin their first clutch earlier than novices, and helpers reduce the time to renest after success from 66 days for an experienced female with no helpers to 50 days for females with at least two years experience and two or more helpers. Breeding females with helpers survive better (76%) than those with no helpers (55%), and helpers thus gain future indirect fitness. Despite their close relatedness to breeders and offspring, in only 19% of group-years did helpers increase their indirect fitness from an increase in productivity.     

Reproductive enhancement by helpers and an experimental inquiry into its mechanism in the bicolored wren

Summary As part of continuing studies of sociality in the wren genus Campylorhynchus we have been studying the bicolored wren — a facultatively cooperative breeder — for the past 6 years in the central Venezuelan savanna. Reproductive groups have ranged in size from 2 to 5. In one of our study populations, only about 15% of the groups contained helpers, and nearly all these contained only a single male helper (Fig. 2). In an adjacent population, the majority of groups contained helpers, and more than half of these contained several helpers of either sex. Territory size is, on average, much smaller in the latter population. In these populations the presence of a single helper is associated with a three-fold increase in reproductive success (Table 1). Additional helpers are not associated with further reproductive enhancement. Enhancement is chiefly due to an increased proportion of nest starts that eventually produce independent juveniles. This reproductive enhancement is not merely an epiphenomenon resulting from the presence of helpers on territories which are superior for other reasons, such as greater resource availability or the quality of particular parents. It is also not a function of the mean or variance in nestling feeding rate. Predator exclusion experiments, in which certan nests were artificially protected from terrestrial predators, suggested that the mechanism of reproductive enhancement was heightened effectiveness of nest defense. Helpers are usually nondispersers from the parental territory, and have always been found to be close relative of the nestlings that they assist in rearing.     

Does the great spotted cuckoo choose magpie hosts according to their parenting ability?

When brood parasites are about to lay an egg, they have to decide which nest to parasitize. The best nest in which to lay will depend on the parenting ability of the host. We have studied selection of magpie (Pica pica) hosts by great spotted cuckoos (Clamator glandarius). Great spotted cuckoos preferentially parasitize large host nests. Nest volume in magpies is a good indicator of territory quality, since there is a negative relationship between magpie nest size and breeding date, and timing of breeding in magpies is known to be positively related to territory quality. Moreover, magpies occupying high-quality territories have high breeding success. Therefore, nest size is positively related to the quality of magpies. Parasitized magpie nests were of greater volume than the nearest neighbouring nest not parasitized by the great spotted cuckoo. In order to test whether the great spotted cuckoos might select high-quality magpie hosts, we manipulated pairs of parasitized and non-parasitized nests with identical laying dates and habitats, introducing into each of the nests the same number of parasitic and non-parasitic eggs. The number of fledglings reared (magpie plus great spotted cuckoo chicks) in naturally parasitized nests was higher than in experimentally parasitized nests. Thus, the probability of survival of the parasite chicks increased if cuckoo eggs were laid in the nests of high-quality hosts originally chosen by the parasite.     

A test of alternate hypotheses for helping behavior in white-fronted bee-eaters of Kenya

Summary Considerable controversy exists over the question of the importance of kin selection in the maintenance of helping behaviors among birds. We examined nine different hypotheses of how helpers might benefit from engaging in alloparental care activities. We break these into four categories: Through its activities the helper may A) improve its probability of surviving to the following breeding season, B) enhance its probability of becoming a breeder in the future, C) increase its reproductive success when it does become a breeder, and D) increase the production of non-descendent kin. The first three categories provide direct fitness gains to the helper; in the fourth, the benefit is indirect. The hypotheses are not mutually exclusive; rather their fitness effects are additive. Each hypothesis, however, makes specific and often separable predictions about both 1) the type of fitness benefits expected, and 2) the characteristics of the birds that serve as helpers. We tested these predictions using five year's data from a color marked and geneologically known population of white-fronted bee-eaters (Merops bullockoides) in Kenya. A) Survival was not related to status (breeder, helper, non-participant); nor did individuals living in large clans have better survival than those living in small ones. B) Newly formed pairs were equally likely to become future breeders irrespective of whether or not one or both individuals had helped previously. C) The mean number of young fledged by a first time breeder was unaffected by its prior helping experience. Neither were first time breeding pairs more likely to gain the services of others as their helpers than were pairs without prior helping experience. Taken together these results demonstrate that beeeaters gain very little direct benefit from alloparenting. D) Helpers did not enhance the survival of the breeders that they helped. But they did have a major effect in increasing nestling survival. Because bee-eater helpers are closely related to the nestlings they help to rear (average r=0.33), they obtain a large indirect benefit by increasing the production of non-descendent kin. We quantified the relative importance of indirect and direct benefits of helping (to the helper) using Vehrencamp's kin index, I _k (1979). I _k compares the fitness consequences of helping against an alternative strategy and calculates the proportion of the inclusive fitness gain or loss that is due to kin (indirect) benefits. Comparing the strategy sets of helping versus not helping for bee-eaters, I _k=0.89 (indicating that 89% of the benefit derived from helping is indirect). When helping was compared against the alternative of breeding, I _k=2.17. Values of I _k greater than 1.0 indicate that direct fitness gains from the alternate strategy (breeding) are greater than those from helping. The value of 2.17 indicates that the helping strategy would not be maintained except for the indirect fitness gained through the increased production of close kin. Alloparenting in white-fronted bee-eaters can thus be considered as altruistic.     

Cooperation in breeding by nonreproductive wrens: kinship,reciprocity, and demography

Summary Stripe-backed wrens (Campylorhynchus nuchalis) often live as adults in large groups on permanent, communally defended territories. Nonbreeding adults cooperate in rearing the young of a single breeding pair; this aid substantially increases the reproductive success of the breeders. In a 6-year study in Venezuela of a completely colorbanded population of 25–30 groups, most adults participated in breeding only as helpers and priority to breeding status was strictly age-determined. Detailed behavioral observations at breeding nests with nestlings showed that, in a sample of 100, helpers nearly always contributed as much to the care of young as breeders. Further, aid-giving does not vary systematically with relatedness of ycung to helpers or with probability of future reciprocation by young. Young being raised are most often at least half siblings of helpers, but seldom return aid to adults that helped raise them. Even adopted helpers collaborate fully. Patterns of demography and dispersal show slow turnover of breeders, delayed reproduction, and a viscous population structure.Application of Hamilton's condition for selection for aid-giving reveals that most individuals in this population can maximize inclusive fitness in the first 2 years by helping instead of breeding. Variation in helping effort and in age of first breeding is related to variation in natal group size and competition resulting from variable demographic neighborhoods in different years or in different parts of the population. Because reciprocation in the form of specific alliance formation among nonreproductives is uncommon, nonspecific reciprocity between cohorts and kin selection account well for the observed pattern of age-dependence in first breeding. Nondiscriminating helping in this population is associated with stable monogamous pair boncs, stable territory boundaries and group membership, strict seniortiy for breeding position, high viscosity and consistent effectiveness of aid. Under these circumstances, very simple behavioral rules amounting to nearly automatic helping seem sufficient to confer critical inclusive fitness gain on helpers.     

Food allocation rules vary with age and experience in a cooperatively breeding parrot

Although it is known that parents can differ in their optimal resource allocation to offspring in size-structured broods, the mechanisms determining differences in allocation rules of carers are not yet clarified. In cooperatively breeding species, breeders and non-reproductive helpers often differ in their fitness payoffs of providing care and in their breeding experience. Cooperative breeders thus provide an appropriate system to examine two hypotheses originally proposed to explain differences in food allocation among parents: (i) food allocation between carers differs because of the distinct cost-benefit ratio of selective feeding (i.e. breeders and helpers are expected to differ in food allocation) and (ii) carers differ in their ability to feed selectively (i.e. differences in food allocation are expected between experienced adults and inexperienced yearlings). We compared feeding rates with which breeders, old helpers and yearling helpers provisioned nestlings of different hatching rank. The influence of experience upon food allocation was further assessed by comparing food allocation of yearlings early and late during nesting. We show that allocation rules differ between age classes because breeders and old helpers fed the youngest chicks most, whereas yearlings showed the opposite pattern. The role of experience was supported by the fact that yearlings adjusted food allocation to that observed in experienced adults during the breeding season. We thus suggest that food allocation in El Oro parakeets depends either on differential skills of adults to transfer food to the youngest chick or on their ability to recognize nestling needs.