Helper effects in the azure‐winged magpie Cyanopica cyana in relation to highly‐clumped nesting pattern and high frequency of conspecific nest‐raiding

In avian cooperative breeding systems, many benefits obtained by social pairs from the presence of helpers have been uncovered. However, until now, the factors that determine the type of assistance helpers provide and the responses of social pairs have not been well illustrated. We examined the contribution of helpers to cooperative groups and the relevant responses of dominant pairs in the azure‐winged magpie Cyanapica cyana which breeds on the Tibetan Plateau. We used the capture–mark–recapture method to identify helpers. Results showed that helpers were mostly the yearling sons of dominant pairs. They mainly contributed to the cooperative group in three ways, courtship‐feeding the incubation female, provisioning the brood, and defending the nest. For responses of dominant pairs, we unexpectedly found that clutch size was not influenced by the presence of helpers at the nest. However, cooperative groups had higher brood feeding rates than biparental nests and their feeding pattern also differed to that of the latter. Consequently, nestlings in cooperative groups had larger fledging body mass than that in biparental nests. By examining reasons for nest failure, we revealed that conspecific nest‐raiding contributed to more nest failure than any other natural predators. Because of the contribution of helpers in defending against both predators and conspecific nest‐raiders, cooperative groups had higher survival rate than biparental nests. Thus, our findings suggest that in a highly‐clumped nesting pattern, factors concerning the risk of nest predation, rather than that influencing food supply, play an important role in determining helper effects and responses of aided dominant pairs.     

The Social System of a Communal Breeder,the Bay-winged Cowbird Molothrus badius

Molothrus badius (bay-winged cowbird), an icterine blackbird with cooperative breeding, shares behavioural and ecological characteristics with other communal nesters: it is sedentary, has a high annual survival rate (76.2%) and a strong nest-site tenacity (mean breeding dispersal of 41.9 and 89.4 m for males and females). Behavioural data, including collective agonistic displays, suggest group territoriality. Before egg hatching most breeders occurred as single pairs showing territorial behaviour (82% of nests), and nesting was usually solitary (distances to nearest nests of 25–103 m). Most breeders were apparently monogamous, with a 2.5% incidence of extrapair copulations in the territory during clutch formation. During the nestling stage one to four helpers occurred at 95% of M. badius nests. Most helpers were 1–2 years old, but older breeding adults (mostly males) that failed to rear their own offspring helped at the end of the season. The number of helpers increased (up to 4) with nestling age. Helpers were also recruited during the postfledging period, and group size reached up to 10 adults at this stage. Helpers mobbed predators and brood parasites, and provided 35% of the nestling food. Provisioning rate was positively and significantly correlated with number of helpers, although age of nestlings was the best predictor of overall food delivery rate. The helping system was almost obligate and productivity comparisons between nests with/without helpers are not possible. Data suggest that helpers increased the breeding success per nest. The correlation between the provisioning rates of parents and helpers was negative but non-significant. In 18% of nests 3 to 4 individuals were present before the nestling period, including cases of apparently polyandrous trios and one case of joint nesting by two pairs. Within Brown 's (1987) categories of social organization M. badius is mainly group territorial with plural nesting. Habitat requirements of M. badius are wide and nest sites do not appear to limit breeding. Kinship plays a role in the social system, as 9 of 12 helpers marked as nestlings helped their parents.     

Helping behaviour in facultatively eusocial hover wasps: an experimental test of the subfertility hypothesis

A candidate explanation for the evolution of eusociality is that helpers are physiologically constrained such that helping is their only realistic option. We tested this subfertility hypothesis in a species of facultatively eusocial hover wasp (Hymenoptera, Stenogastrinae: Liostenogaster flavolineata) by seeing whether helpers that were forced to nest on their own were able to mature their own eggs. One focal helper was left alone on each of 22 nests, from which all other adult wasps (including the dominant) were permanently removed. After 18 days, all but one of the 19 focal helpers that remained on their nests had ovarian development and insemination status characteristic of dominants, and the majority had probably laid eggs. This was in striking contrast to the reproductive status of other helpers removed from the same nests at the start of the experiment. These results provide convincing experimental evidence that females do not become helpers because of some unconditional physiological constraint. There is currently no unequivocal support for the subfertility hypothesis in facultatively eusocial Hymenoptera lacking morphological castes. Copyright 1999 The Association for the Study of Animal Behaviour.     

Fitness consequences of helping behavior in the western bluebird

We examined the fitness consequences of helping behavior inthe western bluebird (Sialia mexicana) at Hastings Reservationin Carmel Valley, California, USA, and tested hypotheses forhow helpers benefit from engaging in alloparental behavior.Both juvenile and adult western bluebirds occasionally helpat the nest During a 12 year period, all adult helpers and mostjuvenile helpers were male. Helpers usually fed at nests ofboth their parents and rarely helped when only one parent waspresent. The frequency of pairs with adult helpers was only7%, but nearly one-third of adult males helped among those withboth parents on the study area. At least 28% were breeders whosenests failed. The propensity to help appears to depend uponparental survival, male philopatry, and the breeding successof potential helpers. Feeding rates were not increased at nestswith juvenile helpers, apparendy because breeding males reducedtheir feeding rates. In contrast, adult helpers increased theoverall rates of food delivery to the nest in spite of a reductionin the number of feeding trips made by both male and femaleparents. Helpers did not derive any obvious direct fitness benefitsfrom helping, but they had greater indirect fitness than nonhelpersdue to increases in nestling growth rates and fledging successat their parents' nests. Helpers fledged fewer offspring intheir first nests than did nonhelpers, suggesting that theywere birds with reduced reproductive potential. Although wehave not yet measured the effect of extrapair fertilizationson the fitness benefits of helping, we calculated the differencein fitness between helpers and nonhelpers as a function of thepotential helper's paternity when breeding independently andhis father's paternity in the nest at which he might help. Inconjunction with constraints on breeding and indirect fitnessbenefits, we predict that relatedness of males to the youngin their own as well as their parents' nests will influencehelping behavior in western bluebirds.     

Helper contributions in the cooperatively breeding laughing kookaburra: feeding young is no laughing matter

I studied the contributions of individuals to incubation and nestling feeding in a population of cooperatively breeding laughing kookaburras, Dacelo novaeguineae. In most cooperatively breeding birds where nest success is limited by nestling starvation, related helpers increase the overall level of provisioning to the nest, thus boosting the production of nondescendent kin. However, although partial brood loss is the largest cause of lost productivity in kookaburra nests, additional helpers failed to increase overall provisioning. Instead, all group members, but especially helpers, reduced their feeding contributions as group size increased. Breeders and helpers reduced the size of prey delivered, and helpers also reduced the number of feeding visits. An important benefit of helping in kookaburras may be to allow all group members to reduce their effort. Within groups, contributions to care depended on status, sex, group size and the brood size. Breeding males delivered the most food. Breeding females provisioned less than their partner, but their effort was comparable to that of male helpers. Female helpers contributed the least food. Incubation effort followed similar patterns. The relatedness of helpers to the brood had no impact on their provisioning. Across all group sizes, helpers generally brought larger items to the nest than breeders. Copyright 2000 The Association for the Study of Animal Behaviour.     

Male Seychelles warblers use territory budding to maximize lifetime fitness in a saturated environment

In cooperatively breeding species, helping at the nest and buddingoff part of the natal territory have been advanced as strategiesto increase fitness in an environment that is saturated withterritories. The importance of helping or territory buddingas a determinant of lifetime reproductive success (LRS) hasbeen debated because the potential benefits of both strategiescould not be separated. Here we test the causes and the immediateand future fitness consequences of single dispersal decisionstaken by male Seychelles warblers (Acrocephalus sechellensis).Males breeding in high-quality territories (high food abundance)have significantly higher LRS than similar-aged males buddingoff part of the parental territory. Initially, budders havea low reproductive success (because of limited food resourcesor absence of a breeding partner). However, they have a longlife span and inherit high-quality territories through sitedominance, by which they gain higher LRS than breeders on low-qualityterritories, helpers, or floaters. Experimental creation ofmale breeding territory vacancies showed that most young malesbecame budders because of intense competition for high-quality territories. The translocation of warblers to the previouslyunoccupied Aride Island shows that males behave according tothe expected fitness benefits of each dispersal strategy. Inthe absence of competition for territories on Aride, all youngmales bred in high-quality territories. However, after saturationof high-quality habitat with territories, most males becamebudders rather than breeders on low-quality habitat, helpers,or floaters.     

Family status and contributions to breeding by Florida scrub jays

Nestling care by Florida scrub jay Aphelocoma c. coerulescens breeders and helpers is quantified. Breeding males and older male helpers deliver more food than first-year helpers and older female helpers. Breeding males adjust the quantity of food they deliver inversely to that delivered by their helpers. Older female helpers, who make many nest visits without delivering food, attempt to usurp the nest-sitting role of the breeding female. The advantages of group breeding to the breeders seem clear. Pairs with helpers live longer and produce more fledglings. The advantages of being a helper, though obscure, probably include non-altruistic factors such as increased survival and, for males, increased chances of obtaining space for breeding. Food contributions of helpers seem to parallel these opportunities.     

A test of the importance of direct and indirect fitness benefits for helping decisions in western bluebirds

In cooperative breeders, the extent to which helpers at thenest adjust their contributions in accordance with direct andindirect (kin-selected) fitness payoffs remains an open question.In a long-term study of the western bluebird, Sialia mexicana,helpers were exclusively male and helped at nests of both parents,a parent and stepparent, or a brother and unrelated female.This natural variation in the context of helping facilitatedcomparison of observational data on groups in which one typeof fitness benefit (current direct, future direct, or indirect)varied, whereas the other two were constant. Helpers reducedtheir share of provisioning as they got older, so comparisonswere restricted to groups with yearling helpers. When potentialdirect fitness benefits were identical, but relatedness wasreduced by half owing to the presence of a stepparent, yearlinghelpers failed to reduce their share of feeding trips to thenest. The potential for future direct fitness benefits via possiblemate and territory inheritance was low, and did not influencethe helper's share of provisioning in a comparison of groupswith similar relatedness and opportunities for current directfitness benefits. Even though cobreeding to gain current directfitness benefits was infrequent (17% of nests with brother-helpers),it was associated with an increase in the helper's share ofprovisioning, suggesting that a helper's feeding allocationresponds positively to increased opportunity for parentage inthe nest. The current study demonstrates a useful frameworkfor separating direct and indirect benefits with respect tohelping decisions, and indicates that western bluebird helpersadjust their feeding rates in response to the potential fordirect fitness benefits in the current nest, not indirect benefitsor future direct fitness payoffs. Although past studies of thispopulation showed that indirect benefits play a role in whetheror not helpers help, the current study indicates that they donot play a role in how frequently helpers feed at the nest.     

The functional significance of multiple nest-building in the Australian Reed Warbler Acrocephalus australis

The vast majority of bird species build a nest in which to breed. Some species build more than one nest, but the function of most multiple nest-building remains unclear. Here we describe the unusual nest-building behaviour of the Australian Reed Warbler Acrocephalus australis , and test experimentally the hypotheses that multiple nest-building is related to individual condition or territory quality, and plays a role in mate assessment. Australian Reed Warblers built two types of nest structures: 'type I' nests, which were used for eggs and nestlings, and 'type II' nests, which were structurally distinct from type I nests, did not support eggs, nestlings or adults and were not essential for successful breeding. The number of type II nests built in each territory varied. Type II nests were only built before breeding had commenced in a territory and females were not observed participating in their construction, supporting a role in female mate choice. Birds provided with supplementary food built significantly more type II nests than control birds. However, supplementary-fed birds did not have greater pairing success, and the addition of further type II nests to territories did not increase the pairing rate or type II nest construction in those territories. There was no relationship between the presence of type II nests and either reproductive success or likelihood of nest predation. We discuss the implications of these results in light of previous suggestions regarding the function of multiple nest-building in birds.     

Positive effects of helpers on reproductive success in the brown treecreeper and the general importance of future benefits

1. Numerous studies of cooperatively breeding species have tested for effects of helpers on reproductive success to evaluate hypotheses for the evolution of cooperation, but relatively few have used experimental or statistical approaches that control for the confounding effects of breeder and territory quality. 2. In the brown treecreeper Climacteris picumnus, most helpers are male offspring of the breeding pair that have delayed dispersal. We analysed 5 years of data (97 territory-years) using hierarchical linear modelling to test for effects of helpers on reproductive success while controlling for confounding factors. 3. The number of helpers was related positively to reproductive success even after controlling for differences between territories and breeders. A threshold effect was observed, with success increasing most with the presence of a second helper (i.e. at group size of four). 4. Feeding at the nest was one mechanism responsible for this effect, as larger groups had higher total feeding rates at all nesting stages. Higher total feeding rates, as well as higher feeding rates by helpers, were correlated in turn with greater reproductive success. 5. An analysis of the effects of helper feeding rate on reproductive success in groups with just one helper produced only weak support for a positive effect of helpers. Controlled comparisons of this kind utilize only a small fraction of the total data available and thus have limited statistical power compared to hierarchical or mixed-modelling. 6. A number of hypotheses to explain the evolution and maintenance of helping behaviour are consistent with our results for brown treecreepers including kin selection and hypotheses based on future direct benefits. 7. A previous synthesis of studies of helper effects that controlled for confounding factors suggested a pattern in which male helpers rarely have positive effects on reproductive success. However, revising that synthesis to include recent hierarchical or mixed-modelling studies suggests that helpers of both sexes usually have positive effects, and that the relative importance of future direct benefits may have been underestimated.     

Flexible Helping Behaviour in the Azure-Winged Magpie

Helping to rear the offspring of others may be a way for younger birds to gain access to future reproduction especially when turnover of breeding opportunities is low. However, this explanation is not applicable to cases where adults also help, or when roles shift between helpers and breeders. Over a period of 6-yr, we studied a marked population of azure-winged Magpie (Cyanopica cyanus) breeding in a non-territorial, colonial system. Magpies bred in a highly flexible cooperative system, in which individuals helped at different stages of the breeding cycle, including nest building, feeding the incubating female and feeding the young and removing the faecal sacs. On average, 50% of hatched nests were assisted by helpers-at-the-nest, and nest success appeared to be positively related to the presence of helpers. Helpers were predominantly males. Although juveniles were more likely to help, both juvenile and adult birds helped. Individual birds behaved as helpers either as a first-option or after having attempted their own breeding (second-option helpers). An individual helper may assist more than one nest during the same breeding season and in different breeding seasons. Reversals between breeder and helper roles were common in both directions, within a breeding season and between years. Helping behaviour is an option for almost any member of the colony. Therefore, hypotheses related to the enhancement of future breeding opportunities for juveniles can be discarded as general explanation of helping in this species. Although the decision to help appeared to be influenced by proximal environmental conditions hindering successful breeding, the associated benefits of helping as opposed to simply recovering for future reproduction, especially for former breeders, deserve further study.     

Behavioural ecology of the North Sulawesi Tarictic Hornbill Penelopides exarhatus exarhatus during the breeding season

Two nests of the North Sulawesi Tarictic Hornbill Penelopides exarhatus exarhatus were monitored for one breeding season (April-July). The females sealed themselves into the nest cavities and remained there for 70–90 days. Incubation was estimated at 16–19 days, and at both nests two offspring fledged. Tarictic Hornbills are cooperative breeders with up to three helpers at the nest and defend foraging territories of 72–139 ha. Food items delivered to the nest included fruits of 34 species (85% of diet items) and invertebrates. Provisioning by helpers allowed breeding males to reduce investment in parental care and may accelerate the development rates in chicks. Constraints on dispersal probably result from habitat saturation rather than living in an unpredictable environment.     

THE PRESENT AND PAST DISTRIBUTION OF THE BALD IBIS IN THE PROVINCE OF THE CAPE OF GOOD HOPE

Tarboton, W. R. 1981. Cooperative breeding and group territoriality in the Black Tit. Ostrich 52:216-225.

In a small, colour-ringed population of Black Tits Parus niger in central Transvaal, 11 of 19 observed breeding units comprised pairs with one to three helper-males. These pairs and groups defended permanent territories, the size of which correlated with the size of the group. There were significantly more territorial disputes during winter when less food was available than in summer. Breeding occurred in summer and the female alone built the nest, incubated the eggs and brooded the young while they were small. During this time she was fed by the alpha male and helper males, although before egg-laying the alpha male prevented helpers from courtship-feeding her. On average, unassisted pairs reared 0,88 young/season whereas pairs with helpers reared 1,55 young/season. However the feeding rate of nestlings of pairs with helpers was not higher than that of unassisted pairs and the number of young reared per group did not correlate with the number of helpers within the group.

The helper system in Black Tits was associated with a skewed sex-ratio (1,7:1 males: females) in the adult population and the data are consistent with the “hopeful reproductive” hypothesis for cooperative breeding.     

Sharing of caring: nestling provisioning behaviour of long-tailed tit, Aegithalos caudatus, parents and helpers

The optimal investment strategies of parents in biparental systems are well studied. This contrasts with a poor theoretical and empirical understanding of variation in individual investment in breeding systems with multiple carers. We used the cooperative breeding system of long-tailed tits, to investigate how parents and helpers adjust their rate of nestling provisioning in relation to measures of nestling demand and the number of helpers. Our aim was to examine whether parents and helpers follow the same provisioning rules. Overall provisioning rates were higher for parents than for helpers. However, both parents and helpers increased their provisioning rates as nestlings aged and provisioned at higher rates early in the day. Parents brought more food to larger broods when not helped, but at nests with helpers, neither parents nor helpers had significantly higher provisioning rates at larger broods. However the total provisioning rate was higher at larger broods at both nests with and without helpers. Parents reduced their work rate in response to the arrival of a helper, but neither parents nor first helpers reduced their work rates further with arrival of additional helpers. Variation in provisioning rates between parents and helpers may be the result of different cost-benefit relations, and a theoretical framework is needed within which to explore the consequences of such differences. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Investment and relatedness: A cost/benefit analysis of breeding and helping in the pied kingfisher (Ceryle rudis)

Helping at the nest in birds is often termed altruism. However, so far, no study has ever demonstrated high costs to a helper's own lifetime reproductive success (=direct fitness), nor its compensation through benefits from relatives other than its own offspring (=indirect fitness). In this paper on pied kingfishers (Ceryle rudis) the relationship between investment, relatedness and inclusive fitness (expressed in terms of genetic equivalents) is investigated for breeding males, and males that help either relatives (=primary helpers) or strangers (=secondary helpers). With respect to guarding nests against predators and feeding young, primary helpers invest as much as breeders, but secondary helpers contribute significantly less. These differences in status and investment (measured in energy expenditure) affect the birds' future to such an extent that primary helpers have a lower chance of surviving and mating than secondary helpers. However, their costs in direct fitness are compensated by pronounced benefits to indirect fitness, resulting from improved survival of siblings and parents. An attempt is made to calculate the inclusive fitness of birds following different strategies over a 2-year period. It is concluded that (a) breeding is superior to helping and helping superior to doing nothing and (b) that kin-selection must be invoked to explain why surplus males choose the more costly primary helper strategy instead of the cheaper secondary helper strategy. Alternative explanations, including group selection, parental manipulation and reciprocity, are discussed.     

Extrapair paternity, inclusive fitness, and within-group benefits of helping in western bluebirds

In central coastal California, USA, 3–16% of western bluebird ( Sialia mexicana ) pairs have adult male helpers at the nest. Demographic data on a colour-ringed population over a 13-year period indicate that helpers gain a small indirect fitness benefit through increases in the number of young fledged from nests of close kin. A small proportion of adult helpers (16%) that were able to breed and help simultaneously had higher annual inclusive fitness than males that only bred. These males comprised such a minor proportion of helpers that the mean fitness of helpers was still lower than the mean fitness of independent breeders. We used DNA fingerprinting to determine whether extrapair fertilizations alter within-group benefits enough to tip the balance in favour of helping behaviour. Overall, 19% of 207 offspring were sired by males other than their social father and extrapair fertilizations occurred in 45% of 51 nests. Intraspecific brood parasitism was rare so that mean mother-nestling relatedness approximated the expected value of 0.5. Extrapair paternity reduced putative father-offspring relatedness to 0.38. Mean helper-nestling relatedness was 0.41 for helpers assisting one or both parents and 0.28 for helpers aiding their brothers. Helpers rarely sired offspring in the nests at which they helped. Helping was not conditional on paternity and helpers were not significantly more closely related to offspring in their parents' nests than to offspring in their own nests. Although helpers may derive extracurricular benefits if helping increases their own or their father's opportunities for extrapair fertilizations, within-nest inclusive fitness benefits of helping do not compensate males for failing to breed. Breeding failure and constraints on breeding are the most likely explanations for why most helpers help.     

Testing common assumptions in studies of songbird nest success

We studied Ovenbird Seiurus aurocapilla and Golden‐winged Warbler Vermivora chrysoptera populations in northern Minnesota, USA, to test two common assumptions in studies of songbird nest success: (1) that the condition of an empty nest on or near its expected fledge date is an indicator of nest fate; and (2) that the presence of a fledgling or family group within a territory confirms a successful nest in that territory. We monitored the condition of nests and used radiotelemetry to monitor juveniles through the expected fledging date and early post‐fledging period. Of nests that contained nestlings 1–2 days before the expected fledge date, fates were misidentified using nest condition alone for 9.5% of Ovenbird nests, but those misidentifications were made in both directions (succeeded or failed), yielding only a small bias in estimated nest success. However, 20% of Golden‐winged Warbler nests were misidentified as successful using nest condition during the final visit interval, biasing the nest success estimate upward by 21–28% depending on the treatment of uncertain nest fates. Fledgling Ovenbirds from 58% of nests travelled beyond their natal territory within 24 h, rising to 98% after 5 days, and those fledglings travelled up to 390 m from nests within 10 days of fledging. Fledgling Golden‐winged Warblers from 13% of nests travelled beyond their natal territory within 24 h, rising to 85% after 5 days, and those fledglings travelled up to 510 m from nests within 10 days of fledging. We conclude that nest condition and fledgling presence can be misleading indicators of nest fate, probably commonly biasing nest success estimates upward, and we recommend that these assumptions should be tested in additional species.     

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.     

Genetic monogamy in single-male groups of acorn woodpeckers, Melanerpes formicivorus

Acorn woodpeckers have one of the most complex social systems of any bird species. Breeding units range in size from monogamous pairs to groups of 15 birds that include multiple breeding males and females as well as nonreproductive helpers-at-the-nest. Groups form when young remain at their natal nest to help their parents breed or when single-sex coalitions of siblings disperse to fill a reproductive vacancy on another territory. Plural breeding and helping behaviour are thought to be favoured through indirect fitness benefits for individuals that would otherwise be unable to breed due to a shortage of reproductive vacancies on territories with acorn stores. We report the results of multi-locus DNA fingerprinting of 51 offspring from 18 nests of 16 socially monogamous pairs of acorn woodpeckers. If socially monogamous females mate outside the pair-bond, indirect fitness benefits for cobreeders and helpers will be significantly reduced. Monogamous pairs accounted for all but one of the 51 offspring we tested; the single exception was apparently sired by the putative father, but the putative mother was excluded from maternity. Our results indicate that individuals remaining on their natal territories as helpers are generally the genetic offspring of the pair they help. They also suggest that single-sex coalitions offspring dispersing together from nests of socially monogamous pairs will be full-siblings.