Helpers and egg investment in the cooperatively breeding acorn woodpecker: testing the concealed helper effects hypothesis.
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ABSTRACT:
SUBMITTER: Koenig WD
PROVIDER: S-EPMC2728902 | biostudies-other | 2009 Sep
REPOSITORIES: biostudies-other
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Project description:Mothers can influence offspring phenotype through egg-mediated maternal effects, which can be influenced by cues mothers obtain from their environment during offspring production. Developing embryos use these components but have mechanisms to alter maternal signals. Here we aimed to understand the role of mothers and embryos in how maternal effects might shape offspring social phenotype. In the cooperatively breeding fish Neolamprologus pulcher different social phenotypes develop in large and small social groups differing in predation risk and social complexity. We manipulated the maternal social environment of N. pulcher females during egg laying by allocating them either to a small or a large social group. We compared egg mass and clutch size and the concentration of corticosteroid metabolites between social environments, and between fertilized and unfertilized eggs to investigate how embryos deal with maternal signalling. Mothers in small groups produced larger clutches but neither laid smaller eggs nor bestowed eggs differently with corticosteroids. Fertilized eggs scored lower on a principal component representing three corticosteroid metabolites, namely 11-deoxycortisol, cortisone, and 11-deoxycorticosterone. We did not detect egg-mediated maternal effects induced by the maternal social environment. We discuss that divergent social phenotypes induced by different group sizes may be triggered by own offspring experience.
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Project description:1. Cooperatively breeding species are typically long lived and hence, according to theory, are expected to maximize their lifetime reproductive success through maximizing survival. Under these circumstances, the presence of helpers could be used to lighten the effort of current reproduction for parents to achieve higher survival. 2. In addition, individuals of different sexes and ages may follow different strategies, but whether male and female breeders and individuals of different ages benefit differently from the presence of helpers has often been overlooked. Moreover, only one study that investigated the relationship between parental survival and the presence of helpers used capture-mark-recapture analyses (CMR). These methods are important since they allow us to account for the non-detection of individuals that are alive in the population but not detected, and thus, the effects on survival and recapture probability to be disentangled. 3. Here, we used multi-event CMR methods to investigate whether the number of helpers was associated with an increase in survival probability for male and female breeders of different ages in the sociable weaver Philetairus socius. In this species, both sexes reduce their feeding rate in the presence of helpers. We therefore predicted that the presence of helpers should increase the breeders' survival in both sexes, especially early in life when individuals potentially have more future breeding opportunities. In addition, sociable weaver females reduce their investment in eggs in the presence of helpers, so we predicted a stronger effect of helpers on female than male survival. 4. As expected we found that females had a higher survival probability when breeding with more helpers. Unexpectedly, however, male survival probability decreased with increasing number of helpers. This antagonistic effect diminished as the breeders grew older. 5. These results illustrate the complexity of fitness costs and benefits underlying cooperative behaviours and how these may vary with the individuals' sex and age. They also highlight the need for further studies on the sex-specific effects of helpers on survival.
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Project description:Among avian cooperative breeders, help in raising offspring is usually provided by males or by both sexes. Sex bias in helping should evolve in response to sex-specific ecological constraints on independent reproduction, with mate shortage for males and breeding vacancy shortage for each sex. Given that male-biased adult sex ratios are prevalent among birds, we predict that male-only helping mainly occurs in temperate species where fast population turnovers deriving from low adult annual survival allow all adult females to hold breeding vacancies, whereas some males overflow as helpers, and both-sex helping in tropical species where saturated habitats prevent not only males, but also females from breeding themselves. As expected, we found that across species, adult survival increased towards tropical zones and warmer climates, and higher adult survival tended to be associated with both-sex helping. Furthermore, sex bias in helping was predicted by latitude and ambient temperature. Our findings of demographic response of species to climate as a potential determinant of bias in helper sex uncover how ecological constraints operate to limit independent reproduction in sex-specific ways.
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Project description: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.
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Project description:In egg laying species, breeding females may adjust the allocation of nutrients or other substances into eggs in order to maximise offspring or maternal fitness. Cooperatively breeding species offer a particularly interesting context in which to study maternal allocation because helpers create predictably improved conditions during offspring development. Some recent studies on cooperative species showed that females assisted by helpers produced smaller eggs, as the additional food brought by the helpers appeared to compensate for this reduction in egg size. However, it remains unclear how common this effect might be. Also currently unknown is whether females change egg composition when assisted by helpers. This effect is predicted by current maternal allocation theory, but has not been previously investigated. We studied egg mass and contents in sociable weavers (Philetairus socius). We found that egg mass decreased with group size, while fledgling mass did not vary, suggesting that helpers may compensate for the reduced investment in eggs. We found no differences in eggs' carotenoid contents, but females assisted by helpers produced eggs with lower hormonal content, specifically testosterone, androstenedione (A4) and corticosterone levels. Taken together, these results suggest that the environment created by helpers can influence maternal allocation and potentially offspring phenotypes.
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Project description: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.
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Project description: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.
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Project description:Phenotypes expressed in a social context are not only a function of the individual, but can also be shaped by the phenotypes of social partners. These social effects may play a major role in the evolution of cooperative breeding if social partners differ in the quality of care they provide and if individual carers adjust their effort in relation to that of other carers. When applying social effects models to wild study systems, it is also important to explore sources of individual plasticity that could masquerade as social effects. We studied offspring provisioning rates of parents and helpers in a wild population of long-tailed tits Aegithalos caudatus using a quantitative genetic framework to identify these social effects and partition them into genetic, permanent environment and current environment components. Controlling for other effects, individuals were consistent in their provisioning effort at a given nest, but adjusted their effort based on who was in their social group, indicating the presence of social effects. However, these social effects differed between years and social contexts, indicating a current environment effect, rather than indicating a genetic or permanent environment effect. While this study reveals the importance of examining environmental and genetic sources of social effects, the framework we present is entirely general, enabling a greater understanding of potentially important social effects within any ecological population.
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Project description:Offspring from elderly parents often have lower survival due to parental senescence. In cooperatively breeding species, where offspring care is shared between breeders and helpers, the alloparental care provided by helpers is predicted to mitigate the impact of parental senescence on offspring provisioning and, subsequently, offspring survival. We test this prediction using data from a long-term study on cooperatively breeding Seychelles warblers (Acrocephalus sechellensis). We find that the nestling provisioning rate of female breeders declines with their age. Further, the total brood provisioning rate and the first-year survival probability of offspring decline progressively with age of the female breeder, but these declines are mitigated when helpers are present. This effect does not arise because individual helpers provide more care in response to the lower provisioning of older dominant females, but because older female breeders have recruited more helpers, thereby receiving more overall care for their brood. We do not find such effects for male breeders. These results indicate that alloparental care can alleviate the fitness costs of senescence for breeders, which suggests an interplay between age and cooperative breeding.
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Project description:We manipulated predation risk in a field experiment with the cooperatively breeding cichlid Neolamprologus pulcher by releasing no predator, a medium- or a large-sized fish predator inside underwater cages enclosing two to three natural groups. We assessed whether helpers changed their helping behaviour, and whether within-group conflict changed, depending on these treatments, testing three hypotheses: 'pay-to-stay' PS, 'risk avoidance' RA, or (future) reproductive benefits RB. We also assessed whether helper food intake was reduced under risk, because this might reduce investments in other behaviours to save energy.Medium and large helpers fed less under predation risk. Despite this effect helpers invested more in territory defence, but not territory maintenance, under the risk of predation (supporting PS). Experimentally covering only the breeding shelter with sand induced more helper digging under predation risk compared to the control treatment (supporting PS). Aggression towards the introduced predator did not differ between the two predator treatments and increased with group member size and group size (supporting PS and RA). Large helpers increased their help ratio (helping effort/breeder aggression received, 'punishment' by the dominant pair in the group) in the predation treatments compared to the control treatment, suggesting they were more willing to PS. Medium helpers did not show such effects. Large helpers also showed a higher submission ratio (submission/ breeder aggression received) in all treatments, compared to the medium helpers (supporting PS).We conclude that predation risk reduces helper food intake, but despite this effect, helpers were more willing to support the breeders, supporting PS. Effects of breeder punishment suggests that PS might be more important for large compared to the medium helpers. Evidence for RA was also detected. Finally, the results were inconsistent with RB.
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