Early and adult social environments have independent effects on individual fitness in a social vertebrate

Evidence that the social environment at critical stages of life-history shapes individual trajectories is accumulating. Previous studies have identified either current or delayed effects of social environments on fitness components, but no study has yet analysed fitness consequences of social environments at different life stages simultaneously. To fill the gap, we use an extensive dataset collected during a 24-year intensive monitoring of a population of Alpine marmots (Marmota marmota), a long-lived social rodent. We test whether the number of helpers in early life and over the dominance tenure length has an impact on litter size at weaning, juvenile survival, longevity and lifetime reproductive success (LRS) of dominant females. Dominant females, who were born into a group containing many helpers and experiencing a high number of accumulated helpers over dominance tenure length showed an increased LRS through an increased longevity. We provide evidence that in a wild vertebrate, both early and adult social environments influence individual fitness, acting additionally and independently. These findings demonstrate that helpers have both short- and long-term effects on dominant female Alpine marmots and that the social environment at the time of birth can play a key role in shaping individual fitness in social vertebrates.     

Helpers increase long-term but not short-term productivity in cooperatively breeding long-tailed tits

Helpers at the nest in the cooperative breeding system of long-tailedtits Aegithalos caudatus exhibit kin preference in their helpingbehavior. The aim of this study was to use multivariate analysesto investigate whether helpers accrue indirect fitness benefitsthrough their cooperation by increasing the productivity ofrelatives. All birds started each season breeding independentlyin pairs, but birds that failed in their own breeding attemptoften redirected their care to help another pair provision theiroffspring. About half of all broods had one or more helpers,86% of which were male. Provisioning rates increased and therewas a corresponding increase in the mass of nestlings withinbroods as the number of helpers increased. Helpers had no significantshort-term effect on productivity because nest predation, nestlingsurvival, and brood size were unaffected by the presence ofhelpers. However, in the long term helpers had a highly significanteffect on the recruitment of fledglings, the positive effectof helpers being linear within the range of helper numbers thatwe observed. We found no evidence to suggest that these resultswere confounded by the effects of individual or habitat quality.We conclude that long-tailed tits accrue indirect fitness benefitsby helping kin. Nevertheless, the inclusive fitness benefitfrom helping is substantially lower than that of independentbreeding, showing that helpers are making the best of a badjob.     

First- and second-order sociality determine survival and reproduction in cooperative cichlids

Cooperative breeders serve as a model to study the evolution of cooperation, where costs and benefits of helping are typically scrutinized at the level of group membership. However, cooperation is often observed in multi-level social organizations involving interactions among individuals at various levels. Here, we argue that a full understanding of the adaptive value of cooperation and the evolution of complex social organization requires identifying the effect of different levels of social organization on direct and indirect fitness components. Our long-term field data show that in the cooperatively breeding, colonial cichlid fish Neolamprologus pulcher, both large group size and high colony density significantly raised group persistence. Neither group size nor density affected survival at the individual level, but they had interactive effects on reproductive output; large group size raised productivity when local population density was low, whereas in contrast, small groups were more productive at high densities. Fitness estimates of individually marked fish revealed indirect fitness benefits associated with staying in large groups. Inclusive fitness, however, was not significantly affected by group size, because the direct fitness component was not increased in larger groups. Together, our findings highlight that the reproductive output of groups may be affected in opposite directions by different levels of sociality, and that complex forms of sociality and costly cooperation may evolve in the absence of large indirect fitness benefits and the influence of kin selection.     

Bi‐parental vs. cooperative breeding in a passerine: fitness‐maximizing strategies of males in response to risk of extra‐pair paternity?

In socially monogamous species, males that risk cuckoldry more than others might gain inclusive fitness by yielding paternity to relatives. We tested this prediction in the Tibetan ground tit Pseudopodoces humilis , an unusual facultative cooperative breeder wherein most helpers (87% males) join a mated pair shortly before clutch completion. Extra-pair paternity among bi-parental broods occurred less often (26% of broods, 9% of young) compared with cooperative broods (68%, 25%). In the former, most extra-pair sires (88%) were pair breeders unrelated to the cuckolded males, whereas in the latter, sires (87%) were mainly helpers related to the dominant males. Brood productivity did not differ between the bi-parental and cooperative breeders, but helpers' partitioning over group paternity reduced the realized reproductive success of helped males. After taking inclusive fitness into account, however, there was no difference in success of dominant males between the two social systems. One possible explanation for the differences in the rates of cuckoldry in the two systems was body size, because pair-bond males in bi-parental situations were significantly larger than those in cooperative ones. We propose two alternative strategies for males to maximize fitness: breed as a pair if large to avoid cuckoldry from helpers, or breed cooperatively if small but compromise some paternity to relatives. Our results provide an unusual route to the incidence of cooperative groups, based on constraints imposed by low competitive ability of breeding males rather than some external ecological or demographic factors.     

Adoption in African wild dogs, Lycaon pictus

Parental care of unrelated young is occasionally misdirected and maladaptive from an individual fitness perspective (e.g. interspecific brood parasitism). Alternatively, adults may'adopt'dependent young and gain direct benefits irrespective of the degree of relatedness. For social species whose fitness is partially a function of group size, direct benefits of increasing group size may provide a mechanism favouring adoption independently of the inclusive fitness consequences. African wild dogs. Lycaon pictus , are social carnivores for which group size has been positively crrelated with reproductive success and competitive ability. Co-operative breeding by wild dogs has been described previously as strongly kin-selected and helpers were assumed to be invariably closely related to offspring. However, observations from this study show that the assumption that helpers are related to those they help is not always valid. A minimum of 25% of study area packs ( n =12) contained nonbreeding adults that provided parental care for unrelated pups. Costs of adopting unrelated weaned wild dog pups are slight or nonexistent except for delayed costs due to later potential conflict with mature same-sex adoptees over access to mates. Immediate and delayed direct benefits identified here are associated with increasing group size and may indicate important mechanisms underlyig adoption and the evolution of helping behaviour in wild dogs.     

Effects of allocare-givers on fitness of infants and parents in callitrichid primates

The effects of callitrichid primate helpers (allocare-givers other than an infant's father) on the survival, reproduction or behavior of infants and parents are reviewed, using both published studies and data from free-ranging golden lion tamarins (Leontopithecus rosalia). Three lines of evidence suggest that helpers may increase their own inclusive fitness: (1) The number of adult males acting as helpers in free-ranging groups is correlated with the number of surviving infants in 3 callitrichid species. However, the lack of a negative correlation with number of infants dying suggests that activities other than direct infant care (e.g. territory defense) may be more important, especially in newly formed groups. (2) In 2 species, captive groups with helpers carry infants for longer periods of time than do groups without helpers. Whether such differences would translate into meaningful survival differences in free-ranging groups is unclear. (3) Helpers reduce the energetic burden of parents by reducing the amount of time they spend transporting or provisioning infants in at least 4 species. Reproductive males are more likely than reproductive females to benefit from the presence of helpers, reducing their investment in infant care activities as the number of helpers in the group increases. In free-ranging golden lion tamarins, the reproductive tenure of males, but not females, increases with the number of helpers in the group, suggesting that a reduction in energetic investment may translate into increased survival. 'Decisions' made by helpers to participate in infant transport are weighed against competing needs for foraging, vigilance, territory defense and, in some cases, prospecting for breeding opportunities. Given this complexity, a sophisticated model may be required to answer the question of how helpers 'decide' to participate in infant care versus other activities.     

Helper effects on pup lifetime fitness in the cooperatively breeding red wolf (Canis rufus)

The evolutionary maintenance of cooperative breeding systems is thought to be a function of relative costs and benefits to breeders, helpers and juveniles. Beneficial effects of helpers on early-life survivorship and performance have been established in several species, but lifetime fitness benefits and/or costs of being helped remain unclear, particularly for long-lived species. We tested for effects of helpers on early- and late-life traits in a population of reintroduced red wolves (Canis rufus), while controlling for ecological variables such as home-range size and population density. We found that the presence of helpers in family groups was positively correlated with pup mass and survival at low population density, but negatively correlated with mass/size at high density, with no relation to survival. Interestingly, mass/size differences persisted into adulthood for both sexes. While the presence of helpers did not advance age at first reproduction for pups of either sex, females appeared to garner long-term fitness benefits from helpers through later age at last reproduction, longer reproductive lifespan and a greater number of lifetime reproductive events, which translated to higher lifetime reproductive success. In contrast, males with helpers exhibited diminished lifetime reproductive performance. Our findings suggest that while helper presence may have beneficial short-term effects in some ecological contexts, it may also incur long-term sex-dependent costs with critical ramifications for lifetime fitness.     

Reproductive success depends on the quality of helpers in the endangered,cooperative El Oro parakeet (Pyrrhura orcesi)

In cooperative species, helping behaviour and reproductive success can be correlated, but understanding this correlation is often impaired by the difficulty to correctly infer causation. While helpers can incur costs by participating in brood care, it is yet unclear if their help depends on their individual quality. We address these questions in the previously unknown cooperative breeding system of the endangered El Oro parakeet (Pyrrhura orcesi). Specifically, we ask (i) whether breeders benefit directly from helpers by an enhanced reproductive success and if so, (ii) whether the amount of this potential benefit is regulated by the quality of contributing group members. Groups consist of a dominant breeding pair accompanied by helpers, but cooperation is not obligate. Microsatellite heterozygosity was used to assess individual quality; its suitability as indicator of quality was reflected in the positive relationship between offspring heterozygosity and recruitment into the population. The reproductive success of breeding pairs depended on helper (genetic) quality and the number of helpers. This relationship occurred on two different levels: clutch size and fledging success, indicating (i) that females profit from high‐quality helpers and probably adjust clutch size accordingly and (ii) that the helpers increase fledging success. Congruently, we found that offspring body condition is positively affected by helper quality, which is most probably explained by the increased feeding rates when helpers are present. We suggest a causal link between cooperation and reproductive success in this frugivorous, endangered parakeet. Further, helper (genetic) quality can be a relevant factor for determining reproductive fitness in cooperative species, particularly in small and bottlenecked populations.     

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.     

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.     

Effects of maternal stress on egg characteristics in a cooperatively breeding fish

Elevated stress experienced by a mother can compromise both her own reproductive success and that of her offspring. In this study, we investigated whether chronically stressed mothers experienced such effects in cooperatively breeding species, in which helpers at the nest potentially compound the negative effects of maternal stress. Using Neolamprologus pulcher, a group-living cichlid fish from Lake Tanganyika, we observed the effects of experimentally increased stress on female reproductive success (measured as inter-spawn interval, and number of eggs) as well as egg characteristics including egg size and cortisol concentrations. Stress levels were manipulated by repeated exposure to the acute stresses of chasing and netting. Stressed females had longer inter-spawn intervals and laid fewer, smaller eggs. Although no significant differences in egg cortisol concentrations were detected between control and stressed females, egg cortisol concentration fell between spawns in control but not in stressed fish. No effect of helper number was detected for any parameter examined, except there appeared to be less change in egg cortisol content in groups with helpers present. Our results suggest that stress imposes fitness costs on breeding females, and social regulation of a dominance hierarchy does not appear to exacerbate or alleviate the negative effects of maternal stress.     

Is there an optimal number of helpers in Alpine marmot family groups?

The consequence of helping behavior on breeders fitness is stillcontroversial. We used multivariate analyses to investigatefor the effects of male and female subordinates on breeders'components of fitness in the Alpine marmot, Marmota marmota.We found that male and female subordinates, respectively, increasedand decreased juvenile survival during winter. Thus, we giveevidence that male subordinates should be considered as helpers,and that helpers provided breeders with immediate reproductivesuccess gains, whereas subordinates females were costly. Helpershad no positive effects on female body condition, on persistence(future survival) of dominants, and on future reproduction (occurrenceand size of a litter). Helpers thus did not provide breederswith delayed fitness benefits, and therfore, the load-lighteninghypothesis was not supported. On the contrary, helpers had delayedfitness cost for dominant males and, consequently, for dominantfemales. Immediate benefits counterbalanced by delayed costssuggested an optimal number of helpers in the family group bothfrom male and female perspectives. An optimality model wellpredicted the observed mean number of helpers in Alpine marmotfamily groups. Optimal numbers of helpers were slightly differentfor males and females, suggesting a potential conflict of interestbetween dominants. We finally discuss the possible mechanismsof helping that may explain the observed pattern in the Alpinemarmot.     

Cooperative Breeding and Group Structure in the Lake Tanganyika Cichlid Neolamprologus savoryi

As yet, cooperative breeding has been described only for some fish species. However, evidence is accumulating that it is widespread among Lake Tanganyika cichlids. We studied the cooperative breeding system of the substrate breeding cichlid Neolamprologus savoryi. Breeding groups typically consisted of a large breeding male with one to four breeding females and three to 33 helpers (mean group size: 14.3 members). Group size was significantly related to breeding male and female body sizes, and larger males had more breeding females and larger sized male helpers. The size of the largest female in the group was positively related to the number and sizes of secondary breeding females and female helpers. In case of multiple breeding females, these females usually divided the group's territory into sub‐territories, each with its own helpers (subgroups). Interspersed between groups, independent fish were detected defending an individual shelter (4.4% of all fish). In 9% of the groups no breeding female was present. All group members participated in territory defence and maintenance, and showed submissive behaviours to larger group members. As expected, the level of between‐subgroup conflicts was high compared with the level of within‐subgroup conflicts. We compare these results with data available from other cooperatively breeding fishes.     

Vigilance and fitness in grey partridges Perdix perdix: the effects of group size and foraging-vigilance trade-offs on predation mortality

1. Vigilance increases fitness by improving predator detection but at the expense of increasing starvation risk. We related variation in vigilance among 122 radio-tagged overwintering grey partridges Perdix perdix (L.) across 20 independent farmland sites in England to predation risk (sparrowhawk Accipiter nisus L., kill rate), use of alternative antipredation behaviours (grouping and use of cover) and survival. 2. Vigilance was significantly higher when individuals fed in smaller groups and in taller vegetation. In the covey period (in early winter when partridges are in flocks), vigilance and use of taller vegetation was significantly higher at sites with higher sparrowhawk predation risk, but tall vegetation was used less by larger groups. Individuals were constrained in reducing individual vigilance by group size and habitat choice because maximum group size was determined by overall density in the area during the covey period and by the formation of pairs at the end of the winter (pair period), when there was also a significant twofold increase in the use of tall cover. 3. Over the whole winter individual survival was higher in larger groups and was lower in the pair period. However, when controlling for group size, mean survival decreased as vigilance increased in the covey period. This result, along with vigilance being higher at sites with increasing with raptor risk, suggests individual vigilance increases arose to reduce short-term predation risk from raptors but led to long-term fitness decreases probably because high individual vigilance increased starvation risk or indicated longer exposure to predation. The effect of raptors on survival was less when there were large groups in open habitats, where individual partridges can probably both detect predators and feed efficiently. 4. Our study suggests that increasing partridge density and modifying habitat to remove the need for high individual vigilance may decrease partridge mortality. It demonstrates the general principle that antipredation behaviours may reduce fitness long-term via their effects on the starvation-predation risk trade-off, even though they decrease predation risk short-term, and that it may be ecological constraints, such as poor habitat (that lead to an antipredation behaviour compromising foraging), that cause mortality, rather than the proximate effect of an antipredation behaviour such as vigilance.     

Breeding success in cooperative meerkats: effects of helper number and maternal state

Studies of cooperatively breeding birds and mammals generallyconcentrate on the effects that helpers have on the number ofreproductive attempts females have per year or on the numberand size of offspring that survive from hatching/weaning toindependence. However, helpers may also influence breeding successbefore hatching or weaning. In the present study, we used anultrasound imager to determine litter sizes close to birth,and multivariate statistics to investigate whether helpers influencefemale fecundity, offspring survival to weaning, and offspringsize at weaning in cooperative meerkats, Suricata suricatta.We found that the number of helpers in a group was correlatedwith the number of litters that females delivered each year,probably because females in large groups gave birth earlierand had shorter interbirth intervals. In addition, althoughpup survival between birth and weaning was primarily influencedby maternal dominance status, helper number may also have asignificant positive effect. By contrast, we found no evidenceto suggest that helpers have a direct effect on either littersizes at birth or pup weights at weaning, which were both significantlyinfluenced by maternal weight at conception. However, becausedifferences in maternal weight were associated with differencesin helper number, helpers have the potential to influence maternalfecundity and offspring size within reproductive attempts indirectly.These results suggest that future studies may need to considerdirect and indirect helper effects on female fecundity and investmentbefore assessing helper effects on reproductive success in societiesof cooperatively breeding vertebrates.     

Paying to stay or paying to breed? Field evidence for direct benefits of helping behavior in a cooperatively breeding fish

Several hypotheses aim to explain the evolution of helping behavior,but conclusive experimental support for evaluating the relativeimportance of individual hypotheses is still lacking. We reporton two field experiments conducted to test the "territory inheritance"and "pay-to-stay" hypotheses in the cooperatively breeding cichlidfish Neolamprologus pulcher The territory inheritance hypothesiswas tested by removing one parent, which created breeding vacancies.In 39% of cases, same-sex helpers took over the breeding spot;in 44% of cases helpers continued helping new breeders, and17% were evicted by new breeders. Helpers that were closelysize matched to the removed breeder had a better chance of gainingthe breeding spot Male helpers tended to continue helping aftera takeover more often than females.The pay-to-stay hypothesiswas tested-by temporarily removing helpers. Whereas breedersdid not respond aggressively to removals, other group membersattacked the removed helpers on their return, and 29% were eventuallyevicted. The returning helpers assisted more by increasing theirrate of territory maintenance and defense and visiting the broodchamber more frequently Size and sex of removed helpers didnot explain the observed aggressive reactions of other groupmembers. Thus, our results support both hypotheses: N. pulcherneeds to pay with help to be allowed to remain protected inthe family group, and there they may inherit the natal territory.N. pulcher helpers gain direct benefits from helping behavior.     

The large-male advantage in brown antechinuses: female choice, male dominance, and delayed male death

Male-biased dimorphism in body size is usually attributed tosexual selection acting on males, through either male competitionor female choice. Brown antechinuses (Antechinus stuartii) aresexually dimorphic in size, and heavier males are known to siremore offspring in the wild. We investigated four possible mechanismsthat might explain this large-male reproductive advantage. Wetested if there is a female preference for large males, a femalepreference for dominant males, if larger males compete moreeffectively for mates, and if there is a survival advantagefor large males during the mating season. We established nestinggroups of males in captivity and conducted mate choice trialsin which males from nesting groups either could or could notinteract. We assessed male dominance rank and recorded survivaltimes after mating. Females did not prefer larger males directly.The results suggest that the other three mechanisms of sexualselection tested account for the large-male advantage: largemales competed more successfully for mates, so were sociallydominant; females rejected subordinates (males they saw losingtwice in contests to previous mates); and dominant males survivedfor longer after their first mating. Females judged male rankbased on direct observation of male competitive interactionsat the time of mating and apparently could not distinguish rankfrom male scent. Effects of size and dominance on male reproductivesuccess are not confounded by age because male antechinusesare semelparous.     

Among-sibling differences in the phenotypes of juvenile fish depend on their location within the egg mass and maternal dominance rank

We investigated whether among-sibling differences in the phenotypes of juvenile fish were systematically related to the position in the egg mass where each individual developed during oogenesis. We sampled eggs from the front, middle and rear thirds of the egg mass in female brown trout of known dominance rank. In the resulting juveniles, we then measured traits that are related to individual fitness: body size, social status and standard metabolic rate (SMR). When controlling for differences among females in mean egg size, siblings from dominant mothers were initially larger (and had a lower mass-corrected SMR) if they developed from eggs at the rear of the egg mass. However, heterogeneity in the size of siblings from different positions in the egg mass diminished in lower-ranking females. Location of the egg within the egg mass also affected the social dominance of the resulting juvenile fish, although the direction of this effect varied with developmental age. This study provides the first evidence of a systematic basis for among-sibling differences in the phenotypes of offspring in a highly fecund organism.     

Vigilance and Collective Detection of Predators in Degus (Octodon degus)

Individuals of social and partially social species typically reduce their vigilance activity when foraging in groups. As a result, per capita risk of predation decreases and individuals allocate more time to foraging and other fitness rewarding activities. Reduction of per capita risk is hypothesized to occur because there are more individuals to detect potential predators. If so, collective (i.e. total) vigilance is expected to increase with foraging group size. Increased surveillance during group foraging may occur if group members scan independently of one another, or sequentially to avoid the overlapping of their vigilance bouts. Intriguingly, such coordinated vigilance assumes that individuals monitor not only the presence, but the vigilance behaviour of group mates. We used seasonal records on time budget and grouping patterns of individually marked degus (Octodon degus), a social rodent, to examine if (a) individual vigilance decreases and/or foraging increases with group size, (b) collective vigilance increases with group size and (c) foraging degus coordinate their vigilance. When foraging, degus decreased their individual vigilance and increased their foraging time when in larger groups. Despite this, degus in larger groups increased their collective vigilance, supporting the hypothesis that socially foraging degus decrease predation risk through an improved ability to detect and escape potential predators. Additionally, patterns of collective vigilance suggested that degus scan independently of each other and so, they do not coordinate their vigilance to prevent its temporal overlapping. This finding does not support that foraging degus monitor the vigilance activity of group mates.     

Cooperative breeding shapes post‐fledging survival in an Afrotropical forest bird

For avian group living to be evolutionary stable, multiple fitness benefits are expected. Yet, the difficulty of tracking fledglings, and thus estimating their survival rates, limits our knowledge on how such benefits may manifest postfledging. We radio‐tagged breeding females of the Afrotropical cooperatively breeding Placid greenbul (Phyllastrephus placidus) during nesting. Tracking these females after fledging permitted us to locate juvenile birds, their parents, and any helpers present and to build individual fledgling resighting datasets without incurring mortality costs or causing premature fledging due to handling or transmitter effects. A Bayesian framework was used to infer age‐specific mortality rates in relation to group size, fledging date, maternal condition, and nestling condition. Postfledging survival was positively related to group size, with fledglings raised in groups with four helpers showing nearly 30% higher survival until independence compared with pair‐only offspring, independent of fledging date, maternal condition or nestling condition. Our results demonstrate the importance of studying the early dependency period just after fledging when assessing presumed benefits of cooperative breeding. While studying small, mobile organisms after they leave the nest remains highly challenging, we argue that the telemetric approach proposed here may be a broadly applicable method to obtain unbiased estimates of postfledging survival.