Egg components and offspring survival vary with group size and laying order in a cooperative breeder.

Prenatal resource allocation to offspring can be influenced by maternal environment and offspring value, and affect offspring survival. An important pathway for flexible maternal allocation is via egg components such as nutrients and hormones. In cooperative breeders, females with helpers may increase resource allocation to eggs-'differential allocation'-or reduce it-'load-lightening'. Yet, helper effects on egg composition have been poorly studied. Moreover, it is unknown how helpers' presence modulates laying order effects on egg content and survival. Here, we investigated how maternal allocation varied with group size and laying order in the cooperatively breeding sociable weaver (Philetairus socius). We estimated interactive effects of helpers and laying order on allocation to egg mass, yolk nutrients-yolk mass, proteins, lipids, carotenoids, vitamin A and vitamin E-and hormones-testosterone, androstenedione, and corticosterone. Results concurred with the 'differential allocation' predictions. Females with more helpers produced later-laid eggs with heavier yolks and more lipids, and laid eggs overall richer in lipids. Proteins, antioxidants, and hormones were not found to vary with helper number. We then analyzed how helper number modulated laying order effects on survival. Females with more helpers did not specifically produce later-laid eggs with higher survival, but eggs laid by females with more helpers were overall more likely to fledge. These findings show that some egg components (yolk mass, lipids) can positively vary according to females' breeding group size, which may improve offspring fitness.

Maternal allocation in relation to weather, predation and social factors in a colonial cooperative bird.

Females may adjust prenatal allocation in relation to ecological conditions that affect reproductive success, such as weather conditions or predation risk. In cooperative breeders, helpers might also influence reproductive success, and previous studies suggest that females can lay smaller eggs or larger clutches when breeding with more helpers. Although recent work suggests that helper effects can vary according to climatic variables, how social and ecological factors interact to shape prenatal allocation is poorly understood. Here, we examine how ecological and social components of the breeding environment covary with egg mass and clutch size, using as a model the sociable weaver Philetairus socius, a colonial, cooperatively breeding passerine. The study spanned 9 years and included over 1,900 eggs from over 550 clutches. Our analyses combined natural variation in weather conditions (rainfall before each reproductive event) with a nest predator-exclusion experiment and continuous monitoring of the mother's social environment, allowing us to estimate how individual females adjust allocation to reproduction as their number of helpers varies. We found that egg mass varied consistently within females and did not clearly differ in relation to rainfall or predation risk. Contrary to previous studies, there was no evidence for plastic adjustments as females gained and lost helpers, and egg mass was instead better predicted by mother size and identity. Females laid larger clutches when breeding in environments where predation risk was experimentally reduced and after higher rainfall levels. Yet, there was no evidence for increasing clutch size as the number of helpers increased, nor for an interaction between helper effects and ecological factors. We conclude that while sociable weaver females can vary their clutch size, they show high individual consistency in egg mass. In addition, we found no evidence that females may maximize fitness through plastic prenatal allocation in relation to the number of helpers, or that the presence/absence of helper effects is modulated by rainfall levels or predation risk. These results challenge our current knowledge on some of the possible benefits of breeding with helpers and call for more long-term analyses on reproductive allocation adjustments in other cooperative systems.

Egg size investment in superb fairy-wrens: helper effects are modulated by climate.

Natural populations might exhibit resilience to changing climatic conditions if they already show adaptive flexibility in their reproductive strategies. In cooperative breeders, theory predicts that mothers with helpers should provide less care when environmental conditions are favourable, but maintain high investment when conditions are challenging. Here, we test for evidence of climate-mediated flexibility in maternal investment in the cooperatively breeding superb fairy-wren Malurus cyaneus We focus on egg size because in this species egg size influences offspring size, and females reduce egg investment when there are helpers at the nest. We report that females lay larger eggs during dry, hot conditions. However, the effect of temperature is modulated by the presence of helpers: the average egg size of females with helpers is reduced during cooler conditions but increased during hot conditions relative to females without helpers. This appears to reflect plasticity in egg investment rather than among female differences. Analysis of maternal survival suggests that helped females are better able to withstand the costs of breeding in hot conditions than females without helpers. Our study suggests that females can use multiple, independent cues to modulate egg investment flexibly in a variable environment.

The role of social environment on parental care: offspring benefit more from the presence of female than male helpers.

Investment in offspring depends on the costs and benefits to the carer, which can vary with sex and social status. Investment also depends on the effort of others by allowing for compensation (load-lightening), with biparental care studies showing that this depends on the state and type of the other carer. By contrast, studies on cooperative breeders have solely focussed on the effects of group size rather than its composition (i.e. social environment). Here we propose and provide the first test of the 'Social Environment' hypothesis, that is, how the characteristics (here the sex) of other helpers present in the group affect parental care and how this in turn affects offspring fitness in cooperatively breeding red-winged fairy-wrens (Malurus elegans). Breeders provisioned nestlings at a higher rate than helpers, but there was no sex difference in provisioning rate. Compensation to increasing group size varied little with sex and status, but strongly depended on social environment. All group members reduced their provisioning rates in response to an increasing number of male (load-lightening), but not female helpers (additive care). As a result, nestlings received more food and grew faster in the presence of female helpers. The increased nestling growth did convey a fitness advantage due to a higher post-fledging survival to adulthood. Our study provides the first evidence that parental care can depend on social environment. This could be an important overlooked aspect to explain variation in parental care in cooperative breeders in general and in particular the enormous variation between the sexes, which we reveal in a literature overview.

Helpers don't help when it's hot in a cooperatively breeding bird, the Southern Pied Babbler.

Cooperative breeding, where more than two individuals invest in rearing a single brood, occurs in many bird species globally and often contributes to improved breeding outcomes. However, high temperatures are associated with poor breeding outcomes in many species, including cooperative species. We used data collected over three austral summer breeding seasons to investigate the contribution that helpers make to daytime incubation in a cooperatively breeding species, the Southern Pied Babbler Turdoides bicolor, and the ways in which their contribution is influenced by temperature. Helpers spent a significantly higher percentage of their time foraging (41.8 ± 13.7%) and a significantly lower percentage of their time incubating (18.5 ± 18.8%) than members of the breeding pair (31.3 ± 11% foraging and 37.4 ± 15.7% incubating). In groups with only one helper, the helper's contribution to incubation was similar to that of breeders. However, helpers in larger groups contributed less to incubation, individually, with some individuals investing no time in incubation on a given observation day. Helpers significantly decrease their investment in incubation on hot days (>35.5°C), while breeders tend to maintain incubation effort as temperatures increase. Our results demonstrate that pied babblers share the workload of incubation unequally between breeders and helpers, and this inequity is more pronounced during hot weather. These results may help to explain why recent studies have found that larger group size does not buffer against the impacts of high temperatures in this and other cooperatively breeding species.

Compensatory and additive helper effects in the cooperatively breeding Seychelles warbler (Acrocephalus sechellensis).

In cooperatively breeding species, care provided by helpers may affect the dominant breeders' investment trade-offs between current and future reproduction. By negatively compensating for such additional care, breeders can reduce costs of reproduction and improve their own chances of survival. Alternatively, helper care can be additive to that of dominants, increasing the fledging fitness of the current brood. However, the influence helpers have on brood care may be affected by group size and territory quality. Therefore, the impact of helping needs to be disentangled from other factors determining offspring investment before conclusive inferences about the effect of help on additive and compensatory care can be made. We used 20 years of provisioning data to investigate the effect of helping on provisioning rates in the facultative cooperatively breeding Seychelles warbler Acrocephalus sechellensis. Our extensive dataset allowed us to statistically disentangle the effects of helper presence, living in larger groups and different food availability. We show compensatory and additive care (i.e., partial compensation) in response to helper provisioning. Helpers lightened the provisioning load of the dominant male and female and increased total provisioning to nestlings. This was irrespective of group size or territory quality (food availability). Moreover, our results illustrate sex-specific variation in parental care over the course of the breeding event. We discriminate between temporal variation, group size, and territory quality processes affecting cooperative care and as such, gain further insight into the importance of these factors to the evolutionary maintenance of helping behavior.

Multiple benefits of alloparental care in a fluctuating environment.

Although cooperatively breeding vertebrates occur disproportionately in unpredictable environments, the underlying mechanism shaping this biogeographic pattern remains unclear. Cooperative breeding may buffer against harsh conditions (hard life hypothesis), or additionally allow for sustained breeding under benign conditions (temporal variability hypothesis). To distinguish between the hard life and temporal variability hypotheses, we investigated whether the number of alloparents at a nest increased reproductive success or load-lightening in superb starlings (Lamprotornis superbus), and whether these two types of benefits varied in harsh and benign years. We found that mothers experienced both types of benefits consistent with the temporal variability hypothesis, as larger contingents of alloparents increased the number of young fledged while simultaneously allowing mothers to reduce their provisioning rates under both harsh and benign rainfall conditions. By contrast, fathers experienced load-lightening only under benign rainfall conditions, suggesting that cooperative breeding may serve to take advantage of unpredictable benign breeding seasons when they do occur. Cooperative breeding in unpredictable environments may thus promote flexibility in offspring care behaviour, which could mitigate variability in the cost of raising young. Our results highlight the importance of considering how offspring care decisions vary among breeding roles and across fluctuating environmental conditions.