The effects of residential locality on parental and alloparental investment among the Aka foragers of the central African Republic

In this paper I examine the intracultural variability of parental and alloparental caregiving among the Aka foragers of the Central African Republic. It has been suggested that maternal kin offer higher frequencies of allocare than paternal kin and that maternal investment in infants will decrease when alloparental assistance is provided. Behavioral observations were conducted on 15 eight- to twelve-monthold infants. The practice of brideservice and the flexibility of Aka residence patterns offered a means to test the effect of maternal residence on parental and alloparental investment. There was significant variation in the frequency of investment and who supplied care to infants depending on whether mothers resided with their kin or their husbands’ kin. However, in spite of the variation in allocare, when all categories of caregivers were examined collectively, infants received similar overall levels of care. Courtney L. Meehan is a Ph.D. student in cultural anthropology at Washington State University. Her research interests include parenting, alloparenting, female social networks, and female-female cooperation and competition.     

The evolution of indicator traits for parental quality: the role of maternal and paternal effects

In systems where individuals provide material resources to their mates or offspring, mate choice based on traits that are phenotypically correlated with the quality of resources provided is expected to be adaptive. Several models have explored the evolution of mating preference where there are direct benefits to choice, but few have addressed how a phenotypic correlation can be established between a male indicator trait and the degree of parental investment. We present a model with three quantitative traits: male and female parental investment and a potential male indicator trait. In our model, the expression of the "indicator" trait in offspring is affected by parental investment. These effects are referred to as maternal or paternal effects, or as "indirect genetic effects" when parental investment is heritable. With genetic variation for degree of parental investment, offspring harbor genes for parental investment that are unexpressed before mating but will affect the investment that they provide when expressed. Because the investment received from the parents affects the expression of the indicator trait, there will be a correlation between the genes for parental investment inherited and the degree of expression of the indicator trait in the offspring. The indicator trait is thus an "honest" signal for the degree of paternal investment.     

Alloparental Care in the African Striped Mouse Rhabdomys pumilio is Age‐Dependent and Influences the Development of Paternal Care

Alloparents contribute to offspring care and alleviate the workload of breeders. The help provided varies with the age and/or experience of helpers, but it is not known whether breeders vary their investment based on the age of helpers by adjusting the parental care they provide. We studied the alloparental care provided by juvenile and subadult philopatric daughters in biparental African striped mice Rhabdomys pumilio with and without the mother as a measure of alleviation of maternal workload. We showed in a previous study that alleviation of maternal workload directly affects the development of paternal care in their sons, so we studied the expression of paternal care in young males raised by helpers as a proxy of the long‐term consequences of helping. Both juvenile and subadult daughters provided care but the level of alloparental care and concomitant alleviation of maternal care was age‐dependent. In the absence of the mother, juvenile daughters provided just 6% of care compared with 24% of subadult daughters. Sons raised by mothers and juvenile helpers displayed the expected exaggerated levels of care also observed when mothers raise litters on their own. While our results show the direct value of subadult daughters, juvenile daughters could contribute indirectly (e.g. nest maintenance) to alleviating maternal workload. The development of paternal care indicates that mothers do distinguish between the care provided by different aged helpers. Overall, the type of alloparental care provided by female striped mice is expected to change over their lifetimes, resulting in increased inclusive fitness through caring for siblings and acquisition of parenting skills.     

The evolution of parental care in the context of sexual selection: a critical reassessment of parental investment theory

Males and females are often defined by differences in their energetic investment in gametes. In most sexual species, females produce few large ova, whereas males produce many tiny sperm. This difference in initial parental investment is presumed to exert a fundamental influence on sex differences in mating and parental behavior, resulting in a taxonomic bias toward parental care in females and away from parental care in males. In this article, we reexamine the logic of this argument as well as the evolutionarily stable strategy (ESS) theory often used to substantiate it. We show that the classic ESS model, which contrasts parental care with offspring desertion, violates the necessary relationship between mean male and female fitness. When the constraint of equal male and female mean fitness is correctly incorporated into the ESS model, its results are congruent with those of evolutionary genetic theory for the evolution of genes with direct and indirect effects. Male parental care evolves whenever half the magnitude of the indirect effect of paternal care on offspring viability exceeds the direct effect of additional mating success gained by desertion. When the converse is true, desertion invades and spreads. In the absence of a genetic correlation between the sexes, the evolution of paternal care is independent of maternal care. Theories based on sex differences in gametic investment make no such specific predictions. We discuss whether inferences about the evolution of sex differences in parental care can hold if the ESS theory on which they are based contains internal contradictions.     

Preferential parental investment in daughters over sons

Female-biased parental investment is unusual but not unknown in human societies. Relevant explanatory models include Fisher’s principle, the Trivers-Willard model, local mate and resource competition and enhancement, and economic rational actor models. Possible evidence of female-biased parental investment includes sex ratios, mortality rates, parents’ stated preferences for offspring of one sex, and direct and indirect measurements of actual parental behavior. Possible examples of female-biased parental investment include the Mukogodo of Kenya, the Ifalukese of Micronesia, the Cheyenne of North America, the Herero of southern Africa, the Kanjar of south Asia, the Mundugumor of New Guinea, contemporary North America, and historical Germany, Portugal, and the United States. Lee Cronk is Assistant Professor of Anthropology at Texas A&M University, College Station, Texas. His main research interests are in human behavioral ecology, reproductive strategies, and East African hunter-gatherers and pastoralists.     

PATERNAL CARE: DIRECT AND INDIRECT GENETIC EFFECTS OF FATHERS ON OFFSPRING PERFORMANCE

Knowledge of how genetic effects arising from parental care influence the evolution of offspring traits comes almost exclusively from studies of maternal care. However, males provide care in some taxa, and often this care differs from females in quality or quantity. If variation in paternal care is genetically based then, like maternal care and maternal effects, paternal effects may have important consequences for the evolution of offspring traits via indirect genetic effects (IGEs). IGEs and direct–indirect genetic covariances associated with parental care can contribute substantially to total heritability and influence predictions about how traits respond to selection. It is unknown, however, if the magnitude and sign of parental effects arising from fathers are the same as those arising from mothers. We used a reciprocal cross‐fostering experiment to quantify environmental and genetic effects of paternal care on offspring performance in the burying beetle, Nicrophorus vespilloides. We found that IGEs were substantial and direct–indirect genetic covariances were negative. Combined, these patterns led to low total heritabilities for offspring performance traits. Thus, under paternal care, offspring performance traits are unlikely to evolve in response to selection, and variation in these traits will be maintained in the population despite potentially strong selection on these traits. These patterns are similar to those generated by maternal care, indicating that the genetic effects of care on offspring performance are independent of the caregiver's sex.     

Should advertising parental care be honest?

Species with paternal care show less exaggerated sexual ornamentation than those in which males do not care, although direct benefits from paternal care can vastly exceed the indirect benefits of mate choice. Whether condition-dependent handicaps can signal parenting ability is controversial. The good-parent process predicts the evolution of honest signals of parental investment, whereas the differential-allocation model suggests a trade-off between the attractiveness of a mate and his care-provisioning. I show that both alternatives can arise from optimal allocations to advertisement, parental investment and future reproductive value of the male, and that the male''s marginal fitness gain from multiple matings determines which option should apply. The marginal gain is diminishing if opportunities for polygyny or extra-pair copulations are limited. Advertisement is then expected to be modest and honest, indicating genetic quality and condition-dependent parental investment simultaneously. Increasing marginal gains are likely to be related to cases where genetic quality has a significant influence on offspring fitness. This alternative leads to differential allocation with stronger advertisement, more frequent extra-pair copulations, and diminished male care. Reliability is also reduced if allocation benefits have thresholds, e.g. if there is a minimum body condition required for survival, or if females use a polygyny-threshold strategy of mate choice.     

Does Absence Matter?

This paper examines the effects of three different types of father absence on the timing of life history events among women in rural Bangladesh. Age at marriage and age at first birth are compared across women who experienced different father presence/absence conditions as children. Survival analyses show that daughters of fathers who divorced their mothers or deserted their families have consistently younger ages at marriage and first birth than other women. In contrast, daughters whose fathers were labor migrants have consistently older ages at marriage and first birth. Daughters whose fathers died when they were children show older ages at marriage and first birth than women with divorced/deserted fathers and women with fathers present. These effects may be mediated by high socioeconomic status and high levels of parental investment among the children of labor migrants, and a combination of low investment, high psychosocial stress, and low alloparental investment among women with divorced/deserted fathers. Our findings are most consistent with the Child Development Theory model of female life history strategies, though the Paternal Investment and Psychosocial Acceleration models also help explain differences between women in low paternal investment situations (e.g., father divorced/abandoned vs. father dead). Father absence in and of itself seems to have little effect on the life history strategies of Bangladeshi women once key reasons for or correlates of absence are controlled, and none of the models is a good predictor of why women with deceased fathers have delayed life histories compared with women whose fathers are present.     

The effect of paternal investment on female fertility intention in South Korea

Life history theory views reproduction as an outcome of resource allocation. The allocation of resources such as parental investments of time, energy and material resources involves trade-offs between number of offspring and timing of reproduction. Within the framework of mammalian parental investment, the outstanding feature of human reproduction is the high level of paternal care. Although empirical evidence suggests that human paternal investment may have evolved as a reproductive strategy to reduce infant and child mortality rates, the effects of actual paternal investment, including allocating time to child care, on female reproductive decisions have received relatively little attention. We examined the trade-off from two perspectives using a representative sample of married South Korean women aged 20–44 in 2005 (n=977). First, paternal investment in domestic labor, including child care and housework, was expected to be associated with women's preference regarding future reproduction. Second, relative paternal investment was expected to increase women's preference for future reproduction, especially among employed women. We found that increased paternal investment in child care and housework remarkably enhanced women's intention to have a second child, especially among employed women. In addition, although family members provide a low percentage of child care in South Korea, such help is likely to be a useful resource for second childbirth among employed women. Somewhat expectedly, older age and longer time since first birth had negative effects on women's second-child intention. There is growing evidence that, in the lowest fertility societies, paternal investment may be an essential resource for promoting future reproductive behavior of women, especially employed women.     

Men’s reproductive investment decisions

Using questionnaire data completed by 170 men, we examine variation in paternal investment in relation to the trade-off between mating and parenting. We found that as men’s self-perceived mate value increases, so does their mating effort, and in turn, as mating effort increases, paternal investment decreases. This study also simultaneously examined the influence on parental investment of men’s mating effort, men’s perception of their mates’ fidelity, and their perceived resemblance to their offspring. All predicted investment. The predictors of investment are also tested independently for men who are still in a relationship with the mother of their children and those that are separated from her. Finally we examine how self-perceived mate value affects how men respond to variation in paternity confidence. Men with a self-perceived low mate value were less likely to respond to lowered mate fidelity by reducing their parental investment compared with men with a self-perceived high mate value.     

The role of maternal and paternal effects in the evolution of parental quality by sexual selection

Genetic models of maternal effects and models of mate choice have focused on the evolutionary effects of variation in parental quality. There have been, however, few attempts to combine these into a single model for the evolution of sexually selected traits. We present a quantitative genetic model that considers how male and female parental quality (together or separately) affect the expression of a sexually selected offspring trait. We allow female choice of males based on this parentally affected trait and examine the evolution of mate choice, parental quality and the indicator trait. Our model reveals a number of consequences of maternal and paternal effects. (1) The force of sexual selection owing to adaptive mate choice can displace parental quality from its natural selection optimum. (2) The force of sexual selection can displace female parental quality from its natural selection optimum even when nonadaptive mate choice occurs (e.g. runaway sexual selection), because females of higher parental quality produce more attractive sons and these sons counterbalance the loss in fitness owing to over-investment in each offspring. (3) Maternal and paternal effects can provide a source of genetic variation for offspring traits, allowing evolution by sexual selection even when those traits do not show direct genetic variation (i.e. are not heritable). (4) The correlation between paternal investment and the offspring trait influenced by the parental effects can result in adaptive mate choice and lead to the elaboration of both female preference and the male sexually selected trait. When parental effects exist, sexual selection can drive the evolution of parental quality when investment increases the attractiveness of offspring, leading to the elaboration of indicator traits and higher than expected levels of parental investment.     

Effects of military activity on breeding birds

United States military bases provide habitat for a diverse suite of wildlife species despite intense anthropogenic disturbance inherent in training activities. Little research has examined how military activity affects wildlife reproduction. We compared parental investment, reproductive success, offspring and adult quality, and stress hormone concentrations of northern cardinals (Cardinalis cardinalis) breeding in areas that differed 10-fold in levels of on-ground military activity. We found no evidence of direct impacts of military activity on cardinals, nor did the reproductive success of several other passerine species appear to be affected. However, we observed American crows (Corvus brachyrhynchos) much less frequently in areas of high military activity, and cardinals nesting in those areas were less responsive to crow models. The apparent displacement of this nest predator suggests that military activity could indirectly benefit some wildlife. Although these results are promising for the conservation of birds that depend on military bases for breeding habitat, further assessment of direct and indirect effects of military training on a wider array of species is required before concluding that such activity is benign. © 2012 The Wildlife Society.     

Paternal-age and birth-order effect on the human secondary sex ratio.

Because of conflicting results in previous analyses of possible maternal and paternal effects on the variation in sex ratio at birth, records of United States live births in 1975 were sorted by offspring sex, live birth order (based on maternal parity), parental races, and, unlike prior studies, ungrouped parental ages. Linear regression and logistic analysis showed significant effects of birth order and paternal age on sex ratio in the white race data (1.67 million births; 10,219 different combinations of independent variables). Contrary to previous reported results, the paternal-age effect cannot be ascribed wholly to the high correlation between paternal age and birth order as maternal age, even more highly correlated with birth order, does not account for a significant additional reduction in sex-ratio variation over that accounted for by birth order alone.     

Establishment of Legal Paternity for Children of Unmarried American Women

The establishment of a legal father for children of unmarried parents reflects both high paternity confidence and male willingness to commit to paternal investment. Whether an unmarried man voluntarily acknowledges paternity after a child is born has important consequences for both the mother and child. This paper brings to bear a life history perspective on paternity establishment, noting that men face trade-offs between mating and parental effort and that women will adjust their investment in children based on expected male investment. I predict that paternity establishment will be more likely when the mother has high socioeconomic status, when maternal health is good, and when the child is male, low parity, or a singleton (versus multiple) birth. I further predict that establishment of paternity will be associated with increased maternal investment in offspring, resulting in healthier babies with higher birthweights who are more likely to be breastfed. These predictions are tested using data on 5.4 million births in the United States from 2009 through 2013. Overall the results are consistent with the hypothesis that the trade-offs men face between reproductive and parental investment influence whether men voluntarily acknowledge paternity when a child is born.     

Relationships of maternal and paternal anthropometry with neonatal body size,proportions and adiposity in an Australian cohort

The patterns of association between maternal or paternal and neonatal phenotype may offer insight into how neonatal characteristics are shaped by evolutionary processes, such as conflicting parental interests in fetal investment and obstetric constraints. Paternal interests are theoretically served by maximizing fetal growth, and maternal interests by managing investment in current and future offspring, but whether paternal and maternal influences act on different components of overall size is unknown. We tested whether parents' prepregnancy height and body mass index (BMI) were related to neonatal anthropometry (birthweight, head circumference, absolute and proportional limb segment and trunk lengths, subcutaneous fat) among 1,041 Australian neonates using stepwise linear regression. Maternal and paternal height and maternal BMI were associated with birthweight. Paternal height related to offspring forearm and lower leg lengths, maternal height and BMI to neonatal head circumference, and maternal BMI to offspring adiposity. Principal components analysis identified three components of variability reflecting neonatal “head and trunk skeletal size,” “adiposity,” and “limb lengths.” Regression analyses of the component scores supported the associations of head and trunk size or adiposity with maternal anthropometry, and limb lengths with paternal anthropometry. Our results suggest that while neonatal fatness reflects environmental conditions (maternal physiology), head circumference and limb and trunk lengths show differing associations with parental anthropometry. These patterns may reflect genetics, parental imprinting and environmental influences in a manner consistent with parental conflicts of interest. Paternal height may relate to neonatal limb length as a means of increasing fetal growth without exacerbating the risk of obstetric complications. Am J Phys Anthropol 156:625–636, 2015. © 2014 The Authors American Journal of Physical Anthropology Published by Wiley Periodicals, Inc.     

The general protected invasion theory: Sex biases in parental and alloparental care

The biases towards eusociality, female workers and maternal care in haplodiploid versus diploid insects may result from the relatively low probabilities that rare mutant, partially dominant alleles promoting these behaviours will be lost by genetic drift in haplodiploid populations (Reeve, 1993). A generalization of this 'protected invasion' theory also predicts that parental and alloparental care will tend to be associated with the homogametic sex in diploid populations if the Y chromosome of the heterogametic sex is absent or largely inert. Sex differences in (allo)parental care (i.e. either parental or alloparental care) should increase with increased asymmetry between the sexes in the fraction of behaviour-influencing loci occurring on their characteristic sex chromosomes. The theory explains the strong predisposition towards female (allo)parental care in mammals, a contrasting tendency towards male (allo)parental care in birds, the propensity for joint male and female (allo)parental care in termites, and biases towards female cooperation in social spiders. The theory also explains the apparent rarity or absence of alloparental care in marsupials, an intriguing consequence of preferential paternal X-chromosome inactivation in this taxon. Thus protected invasion theory possibly provides new insights into the relationship between social structure and the genetic system. The theory does not compete with ecological or kin-selective hypotheses for the advantages of (allo)parental care; indeed, such advantages must exist for protected-invasion biases to operate.     

Behavioural and morphometric measurements of parental investment in southern elephant seals at the Falkland Islands

Parental investment is a key variable in the study of breeding strategies and life-histories evolution. In Pinnipedia, parental investment is usually calculated from direct measurements of pup weight gain or energy transfer between the mother and the pup. These direct methods always involve handling and restraining procedures that pose practical, logistical and ethical problems. To evaluate if weighing can be substituted by indirect observational estimates of parental investment, we analysed the relationship among various behavioural measures of suckling and post-natal growth in the southern elephant seal population of Sea Lion Island (Falkland Islands). Behavioural measures were in all cases a poor predictor of true investment as estimated by weighing. We concluded that there are currently no effective alternatives to direct handling, and that the best way to reduce the potential adverse impact of investment studies is the improvement of the handling protocol, which should include an estimation of the long-term effects on the health of handled animals. Further research is needed to test the validity of non-behavioural indirect methods (e.g. 3D photogrammetry).     

A model for evolution of male parental care and female multiple mating

In most animals, males gain a fitness benefit by mating with many females, whereas the number of progeny per female is unlikely to increase as a function of additional mates. Furthermore, males of internally fertilizing species run the risk of investing in offspring of other males if they provide parental care. Nevertheless, males of many avian species and a minority of mammalian species provide parental care, and females of various species mate with multiple males. I investigate a two-locus genetic model for evolution of male parental care and female multiple mating in which females gain a direct benefit by multiple mating from the paternal care they thereby elicit for their offspring. The model suggests that, first, male parental care can evolve when it strongly enhances offspring survival and the direct costs of female multiple mating (e.g., loss of energy, risk of injury, exposure to infectious diseases) are greater than its indirect benefit (e.g., acquisition of good genes, increased genetic diversity among offspring); second, female multiple mating can evolve when paternal care is important for offspring survival or the indirect benefit of multiple mating is larger than its direct cost; and, finally, male parental care and female multiple mating can co-occur.     

A comparison of paternal behaviour in the meadow vole Microtus pennsylvanicus,the pine vole M. pinetorum and the prairie vole M. cchrogaster

Paternal care in microtines has been studied infrequently and few studies have compared patterns of direct and indirect paternal investment. The paternal behaviour of three vole species, the meadow vole (Microtus pennsylvanicus), the pine vole (M. pinetorum) and the prairie vole (M. ochrogaster) was examined in a semi-natural setting. Prairie and pine voles were found to exhibit high levels of paternal care. Prairie vole males contributed the most direct care by remaining in the natal nest for long periods of time in contact with the pups. Pine voles contributed less direct care than prairie voles as they spent less time in the natal nest with their offspring. In addition, both prairie and pine vole males were observed to groom their pups and retrieve them back to the nest area. Prairie vole males also engaged in such indirect forms of care as nest construction and maintenance, while pine voles provided indirect care in the form of tunnel construction and food caching. Meadow vole males were the least paternal of the three species and rarely engaged in either direct or indirect care. These findings support predictions that M. pennsylvanicus is promiscuous and that male and female meadow voles occupy separate territories. They are also consistent with studies which indicate that prairie and pine voles are monogamous and have a structured social organization with members interacting closely with one another.     

Kinship affects investment by helpers in a cooperatively breeding bird

Helping behaviour in cooperative breeding systems has been attributed to kin selection, but the relative roles of direct and indirect fitness benefits in the evolution of such systems remain a matter of debate. In theory, helpers could maximize the indirect fitness benefits of cooperation by investing more in broods with whom they are more closely related, but there is little evidence for such fine-scale adjustment in helper effort among cooperative vertebrates. In this study, we used the unusual cooperative breeding system of the long-tailed tit Aegithalos caudatus to test the hypothesis that the provisioning effort of helpers was positively correlated with their kinship to broods. We first use pedigrees and microsatellite genotypes to characterize the relatedness between helpers and breeders from a 14 year field study. We used both pedigree and genetic approaches because long-tailed tits have access to pedigree information acquired through social relationships, but any fitness consequences will be determined by genetic relatedness. We then show using both pedigrees and genetic relatedness estimates that alloparental investment by helpers increases as their relatedness to the recipients of their care increases. We conclude that kin selection has played a critical role in moulding the investment decisions of helpers in this cooperatively breeding species.