Investment by maternal grandmother buffers children against the impacts of adverse early life experiences.

Exogenous shocks during sensitive periods of development can have long-lasting effects on adult phenotypes including behavior, survival and reproduction. Cooperative breeding, such as grandparental care in humans and some other mammal species, is believed to have evolved partly in order to cope with challenging environments. Nevertheless, studies addressing whether grandparental investment can buffer the development of grandchildren from multiple adversities early in life are few and have provided mixed results, perhaps owing to difficulties drawing causal inferences from non-experimental data. Using population-based data of English and Welsh adolescents (sample size ranging from 817 to 1197), we examined whether grandparental investment reduces emotional and behavioral problems in children resulting from facing multiple adverse early life experiences (AELEs), by employing instrumental variable regression in a Bayesian structural equation modeling framework to better justify causal interpretations of the results. When children had faced multiple AELEs, the investment of maternal grandmothers reduced, but could not fully erase, their emotional and behavioral problems. No such result was observed in the case of the investment of other grandparent types. These findings indicate that in adverse environmental conditions the investment of maternal grandmothers can improve child wellbeing.

Parental Investment by Birth Fathers and Stepfathers : Roles of Mating Effort and Childhood Co-residence Duration.

This study investigates the determinants of paternal investment by birth fathers and stepfathers. Inclusive fitness theory predicts higher parental investment in birth children than stepchildren, and this has consistently been found in previous studies. Here we investigate whether paternal investment varies with childhood co-residence duration and differs between stepfathers and divorced birth fathers by comparing the investment of (1) stepfathers, (2) birth fathers who are separated from the child's mother, and (3) birth fathers who still are in a relationship with her. Path analysis was conducted using cross-sectional data from adolescents and younger adults (aged 17-19, 27-29, and 37-39 years) from the German Family Panel (pairfam), collected in 2010-2011 (n = 8326). As proxies of paternal investment, we used financial and practical help, emotional support, intimacy, and emotional closeness, as reported by the children. We found that birth fathers who were still in a relationship with the mother invested the most, and stepfathers invested the least. Furthermore, the investment of both separated fathers and stepfathers increased with the duration of co-residence with the child. However, in the case of financial help and intimacy, the effect of childhood co-residence duration was stronger in stepfathers than in separated fathers. Our findings support inclusive fitness theory and mating effort theory in explaining social behavior and family dynamics in this population. Furthermore, social environment, such as childhood co-residence was associated with paternal investment.

Differential grandparental investment when maternal grandmothers are living versus deceased.

Grandparents can increase their inclusive fitness by investing time and resources in their grandchildren. However, not all grandparents make such investments equally, and between-grandparent differences in this regard can be predicted based on paternity uncertainty, lineage and grandparents' sex. Using population-based data for English and Welsh adolescents (n = 1430), we examined whether the death of the most important grandparent (in terms of investment), the maternal grandmother (MGM), changes relative support for existing hypotheses predicting differential grandparental-investment patterns. To contrast the predictions of the grandparental investment hypotheses, we used generalized order-restricted information criterion approximation. We consequently found that, when MGMs are alive, the most-supported hypothesis is 'discriminative grandparental solicitude', which ranks grandparental investment as MGMs > maternal grandfathers (MGFs) > paternal grandmothers (PGMs) > paternal grandfathers (PGFs). However, when MGMs are deceased, the paternity uncertainty hypothesis (MGFs = PGMs > PGFs) receives the most support; this is due to increased investment by PGMs. Thus, when the heaviest investors (i.e. MGMs) are deceased, PGM investments are closer to-but do not exceed-MGF investments.

The interplay of grandparental investment according to the survival status of other grandparent types.

Inclusive fitness theory predicts that grandparental investment in grandchildren aims to maximise their inclusive fitness. Owing to an increasing overlap between successive generations in modern affluent populations, the importance of grandparental investment remains high. Despite the growing literature, there is limited knowledge regarding how the survival status of different grandparent types influences each other's investment in grandchildren. This question was studied by using the Involved Grandparenting and Child Well-Being Survey, which provided nationally representative data of English and Welsh adolescents aged 11-16-years. We applied Bayesian structural equation modeling (BSEM) where grandparental investment in grandchildren was modelled using multi-indicator unobserved latent variable. Our results showed that maternal grandmothers' investment was increased by having a living maternal grandfather but not vice versa. Having a living maternal grandmother was also associated with decreased investment of paternal grandparents while the opposite was not found. These findings indicate that the association between the survival status of other grandparents and the focal grandparents' investment varies between grandparent types.

Mothers with higher twinning propensity had lower fertility in pre-industrial Europe.

Historically, mothers producing twins gave birth, on average, more often than non-twinners. This observation has been interpreted as twinners having higher intrinsic fertility - a tendency to conceive easily irrespective of age and other factors - which has shaped both hypotheses about why twinning persists and varies across populations, and the design of medical studies on female fertility. Here we show in >20k pre-industrial European mothers that this interpretation results from an ecological fallacy: twinners had more births not due to higher intrinsic fertility, but because mothers that gave birth more accumulated more opportunities to produce twins. Controlling for variation in the exposure to the risk of twinning reveals that mothers with higher twinning propensity - a physiological predisposition to producing twins - had fewer births, and when twin mortality was high, fewer offspring reaching adulthood. Twinning rates may thus be driven by variation in its mortality costs, rather than variation in intrinsic fertility.

Structural equation modeling reveals decoupling of ecological and self-perceived outcomes in a garden box social-ecological system.

It is well known that green urban commons enhance mental and physical well-being and improve local biodiversity. We aim to investigate how these outcomes are related in an urban system and which variables are associated with better outcomes. We model the outcomes of an urban common-box gardening-by applying the Social-Ecological Systems (SES) framework. We expand the SES framework by analyzing it from the perspective of social evolution theory. The system was studied empirically through field inventories and questionnaires and modeled quantitatively by Structural Equation Modeling (SEM). This method offers powerful statistical models of complex social-ecological systems. Our results show that objectively evaluated ecological outcomes and self-perceived outcomes are decoupled: gardening groups that successfully govern the natural resource ecologically do not necessarily report many social, ecological, or individual benefits, and vice versa. Social capital, box location, gardener concerns, and starting year influenced the changes in the outcomes. In addition, the positive association of frequent interactions with higher self-perceived outcomes, and lack of such association with relatedness of group members suggests that reciprocity rather than kin selection explains cooperation. Our findings exemplify the importance of understanding natural resource systems at a very low "grassroot" level.

Matrilateral bias of grandparental investment in grandchildren persists despite the grandchildren's adverse early life experiences.

Evolutionary theory predicts a downward flow of investment from older to younger generations, representing individual efforts to maximize inclusive fitness. Maternal grandparents and maternal grandmothers (MGMs) in particular consistently show the highest levels of investment (e.g. time, care and resources) in their grandchildren. Grandparental investment overall may depend on social and environmental conditions that affect the development of children and modify the benefits and costs of investment. Currently, the responses of grandparents to adverse early life experiences (AELEs) in their grandchildren are assessed from a perspective of increased investment to meet increased need. Here, we formulate an alternative prediction that AELEs may be associated with reduced grandparental investment, as they can reduce the reproductive value of the grandchildren. Moreover, we predicted that paternal grandparents react more strongly to AELEs compared to maternal grandparents because maternal kin should expend extra effort to invest in their descendants. Using population-based survey data for English and Welsh adolescents, we found evidence that the investment of maternal grandparents (MGMs in particular) in their grandchildren was unrelated to the grandchildren's AELEs, while paternal grandparents invested less in grandchildren who had experienced more AELEs. These findings seemed robust to measurement errors in AELEs and confounding due to omitted shared causes.

Identifying the paths of climate effects on population dynamics: dynamic and multilevel structural equation model around the annual cycle.

How environmental factors influence population dynamics in long-distance migrants is complicated by the spatiotemporal diversity of the environment the individuals experience during the annual cycle. The effects of weather on several different aspects of life history have been well studied, but a better understanding is needed on how weather affects population dynamics through the different associated traits. We utilise 77 years of data from pied flycatcher (Ficedula hypoleuca), to identify the most relevant climate signals associated with population growth rate. The strongest signals on population growth were observed from climate during periods when the birds were not present in the focal location. The population decline was associated with increasing precipitation in the African non-breeding quarters in the autumn (near the arrival of migrants) and with increasing winter temperature along the migration route (before migration). The number of fledglings was associated positively with increasing winter temperature in non-breeding area and negatively with increasing winter temperature in Europe. These possible carry-over effects did not arise via timing of breeding or clutch size but the exact mechanism remains to be revealed in future studies. High population density and low fledgling production were the intrinsic factors reducing the breeding population. We conclude that weather during all seasons has the potential to affect the reproductive success or population growth rate of this species. Our results show how weather can influence the population dynamics of a migratory species through multiple pathways, even at times of the annual cycle when the birds are in a different location than the climate signal.

Impact of continuous predator threat on telomere dynamics in parent and nestling pied flycatchers.

In addition to direct mortality, predators can have indirect effects on prey populations by affecting prey behaviour or physiology. For example, predator presence can increase stress hormone levels, which can have physiological costs. Stress exposure accelerates the shortening of telomeres (i.e. the protective caps of chromosomes) and shorter telomeres have been linked to increased mortality risk. However, the effect of perceived predation risk on telomeres is not known. We investigated the effects of continuous predator threat (nesting Eurasian pygmy owl Glaucidium passerinum) on telomere dynamics of both adult and partially cross-fostered nestling pied flycatchers (Ficedula hypoleuca) in the wild. Females nesting at owl-inhabited sites showed impaired telomere maintenance between incubation and chick rearing compared to controls, and both males and females ended up with shorter telomeres at owl-inhabited sites in the end of chick rearing. On the contrary, both original and cross-fostered chicks reared in owl sites had consistently longer telomeres during growth than chicks reared at control sites. Thus, predation risk may cause a long-term cost in terms of telomeres for parents but not for their offspring. Predators may therefore affect telomere dynamics of their preys, which could have implications for their ageing rate and consequently for population dynamics.

Does testosterone predict women's preference for facial masculinity?

The influence of sex hormones on women's mate preferences has been an intensively discussed topic for more than a decade. Yet the extent to which levels of sex hormones, and testosterone in particular, influence women's mate preferences is unclear. Thus, the current study used multilevel modelling to investigate putative relationships between salivary testosterone and facial masculinity preferences in a sample of 68 women, while controlling for their age, partnership status, and sociosexuality. We found no significant associations between masculinity preferences and either individual differences or within-woman changes in testosterone. We did find however, that sociosexuality was positively correlated with masculinity preferences. Although it has previously been suggested that testosterone is related to women's facial masculinity preference, our data do not support this proposal.

Males have more aggressive and less sociable personalities than females in semi-captive Asian elephants.

Personality, i.e. consistent between-individual differences in behaviour, has been documented in many species. Yet little is known about how males and females of long-lived, highly social species differ in their measures of personality structure. We investigated sex differences in the mean, variance, and covariance of three previously reported personality traits (Attentiveness, Sociability, Aggressiveness) in 150 female and 107 male Asian elephants (Elephas maximus) from a semi-captive population in Myanmar. These three personality traits were obtained by performing exploratory factor analysis on 28 behavioural items that had been rated by experienced elephant handlers. We found that males scored significantly higher on Aggressiveness and tended to score lower on Sociability than females. However, no sex difference was found in the mean scores of Attentiveness. Variances for the three personality traits did not differ between the sexes, suggesting that male and female elephants share the same range of personality variation. Likewise, trait covariances were similar between the sexes. While both sexes show complex sociality in the wild, female Asian elephants typically live in highly social family units, whereas male elephants' social bonds are weaker. Males usually form dominance ranks by aggressive interactions, especially during musth. Our results on a large sample of individuals living in their natural environment are thus in agreement with elephant life-histories and parallel the findings of sex differences in other long-lived highly social species with similar life-histories.

Search for a Resource-Based Trade-off Between Lifetime Reproductive Effort and Women's Postreproductive Survival in Preindustrial Sweden.

A reduced survival as a potential cost of high lifetime reproductive effort in women has intrigued human evolutionary biologists for more than a century. However, we do not currently have compelling evidence for the delayed survival costs of reproduction. Reasons for this may include several methodological issues, such as environmental confounding, measurement of individuals' lifetime reproductive effort using demographic data, and the practice of mortality selection that are all likely to compromise our ability to reliably detect trade-offs at the phenotypic level. The current research aims to address all these issues by using structural equation modeling to examine the potential trade-off between women's lifetime reproductive effort and their postreproductive mortality in a large data set of 6,594 women from preindustrial northern Sweden that has not previously been used for this purpose. Despite this, the results showed only weak evidence for a trade-off between lifetime reproductive effort and postreproductive mortality, one that was confined to only those women who had high lifetime reproductive effort and spend more than 25 years in widowhood. The socioeconomic status of the family or mother's ethnic background did not moderate this association, with the general trend being one of higher, not lower, postreproductive survival with high lifetime reproductive effort in women.

Evaluating the personality structure of semi-captive Asian elephants living in their natural habitat.

Data on personality for long-lived, highly social wild mammals with high cognitive abilities are rare. We investigated the personality structure of Asian elephants (Elephas maximus) by using a large sample of semi-captive timber elephants in Myanmar. Data were collected during 2014-2017 using questionnaires, for which elephant riders (mahouts) rated 28 behavioural adjectives of elephants. Repeated questionnaires were obtained for each elephant from several raters whenever possible, resulting in 690 ratings of 150 female and 107 male elephants. We started by performing a confirmatory factor analysis to compare the fit of our data to a previously published captive elephant personality structure. Owing to a poor fit of this model to our data, we proceeded by performing explanatory factor analysis to determine the personality structure in our study population. This model suggested that personality in these elephants was manifested as three factors that we labelled as Attentiveness, Sociability and Aggressiveness. This structure did not differ between the sexes. These results provide the basis for future research on the link between personality and reproductive success in this endangered species and more generally, help to resolve the selective pressures on personalities in long-lived, highly social species.

Accounting for measurement error in human life history trade-offs using structural equation modeling.

OBJECTIVES: Revealing causal effects from correlative data is very challenging and a contemporary problem in human life history research owing to the lack of experimental approach. Problems with causal inference arising from measurement error in independent variables, whether related either to inaccurate measurement technique or validity of measurements, seem not well-known in this field. The aim of this study is to show how structural equation modeling (SEM) with latent variables can be applied to account for measurement error in independent variables when the researcher has recorded several indicators of a hypothesized latent construct. METHODS: As a simple example of this approach, measurement error in lifetime allocation of resources to reproduction in Finnish preindustrial women is modelled in the context of the survival cost of reproduction. In humans, lifetime energetic resources allocated in reproduction are almost impossible to quantify with precision and, thus, typically used measures of lifetime reproductive effort (e.g., lifetime reproductive success and parity) are likely to be plagued by measurement error. These results are contrasted with those obtained from a traditional regression approach where the single best proxy of lifetime reproductive effort available in the data is used for inference. RESULTS: As expected, the inability to account for measurement error in women's lifetime reproductive effort resulted in the underestimation of its underlying effect size on post-reproductive survival. CONCLUSIONS: This article emphasizes the advantages that the SEM framework can provide in handling measurement error via multiple-indicator latent variables in human life history studies.

Selection bias in studies of human reproduction-longevity trade-offs.

A shorter lifespan as a potential cost of high reproductive effort in humans has intrigued researchers for more than a century. However, the results have been inconclusive so far and despite strong theoretical expectations we do not currently have compelling evidence for the longevity costs of reproduction. Using Monte Carlo simulation, it is shown here that a common practice in human reproduction-longevity studies using historical data (the most relevant data sources for this question), the omission of women who died prior to menopausal age from the analysis, results in severe underestimation of the potential underlying trade-off between reproduction and lifespan. In other words, assuming that such a trade-off is expressed also during reproductive years, the strength of the trade-off between reproduction and lifespan is progressively weakened when women dying during reproductive ages are sequentially and non-randomly excluded from the analysis. In cases of small sample sizes (e.g. few hundreds of observations), this selection bias by reducing statistical power may even partly explain the null results commonly found in this field. Future studies in this field should thus apply statistical approaches that account for or avoid selection bias in order to recover reliable effect size estimates between reproduction and longevity.

Human reproduction and health: an evolutionary perspective.

According to life history theory, increased investment in reproductive function (physiology and behaviour) at different times throughout the life course affects the risk of many diseases and, ultimately, longevity. Although genetic factors contribute to interindividual and interpopulation variation in reproductive traits, the dominant source of variability is phenotypic plasticity during development and adult life. Reproductive traits in both sexes evolved sensitivity to ecological conditions, as reflected in contemporary associations of hormone concentrations with geographical setting, nutritional status, and physical activity level. Lifetime exposure to increased concentrations of sex hormones is associated with the risk of some cancers, hence decreasing fertility patterns contribute to secular increases in their incidence. Conversely, increased investment in reproductive function might compromise somatic investment in health, such that faster sexual maturation and higher parity increases risk of diabetes and cardiovascular disease. An evolutionary perspective on reproductive biology could improve the efficacy of public health efforts to reduce the risk of hormone-sensitive cancers and other non-communicable diseases.

Using change trajectories to study the impacts of multi-annual habitat loss on fledgling production in an old forest specialist bird.

The loss and subdivision of habitat into smaller and more spatially isolated units due to human actions has been shown to adversely affect species worldwide. We examined how changes in old forest cover during eight years were associated with the cumulative number of fledged offspring at the end of study period in Eurasian treecreepers (Certhia familiaris) in Central Finland. We were specifically interested in whether the initial level of old forest cover moderated this relation. We applied a flexible and powerful approach, latent growth curve modelling in a structural equation modeling (SEM) framework, to create trajectories describing changes in old forest cover through time, and studied how this change at both the territory core and landscape scales impacted fledging numbers. Our main finding was that at the territory core scale the negative impact of habitat loss on fledging numbers was lessened by the higher levels of initial forest cover, while no association was found at the landscape scale. Our study highlights a powerful, but currently under-utilised methodology among ecologists that can provide important information about biological responses to changes in the environment, providing a mechanistic way to study how land cover dynamics can affect species responses.

Natural selection for earlier male arrival to breeding grounds through direct and indirect effects in a migratory songbird.

For migratory birds, the earlier arrival of males to breeding grounds is often expected to have fitness benefits. However, the selection differential on male arrival time has rarely been decomposed into the direct effect of male arrival and potential indirect effects through female traits. We measured the directional selection differential on male arrival time in the pied flycatcher (Ficedula hypoleuca) using data from 6 years and annual number of fledglings as the fitness proxy. Using structural equation modeling, we were able to take into account the temporal structure of the breeding cycle and the hierarchy between the examined traits. We found directional selection differentials for earlier male arrival date and earlier female laying date, as well as strong selection differential for larger clutch size. These selection differentials were due to direct selection only as indirect selection for these traits was nonsignificant. When decomposing the direct selection for earlier male arrival into direct and indirect effects, we discovered that it was almost exclusively due to the direct effect of male arrival date on fitness and not due to its indirect effects via female traits. In other words, we showed for the first time that there is a direct effect of male arrival date on fitness while accounting for those effects that are mediated by effects of the social partner. Our study thus indicates that natural selection directly favored earlier male arrival in this flycatcher population.

Culled males, infant mortality and reproductive success in a pre-industrial Finnish population.

Theoretical and empirical literature asserts that the sex ratio (i.e. M/F) at birth gauges the strength of selection in utero and cohort quality of males that survive to birth. We report the first individual-level test in humans, using detailed life-history data, of the 'culled cohort' hypothesis that males born to low annual sex ratio cohorts show lower than expected infant mortality and greater than expected lifetime reproductive success. We applied time-series and structural equation methods to a unique multigenerational dataset of a natural fertility population in nineteenth century Finland. We find that, consistent with culled cohorts, a 1 s.d. decline in the annual cohort sex ratio precedes an 8% decrease in the risk of male infant mortality. Males born to lower cohort sex ratios also successfully raised 4% more offspring to reproductive age than did males born to higher cohort sex ratios. The offspring result, however, falls just outside conventional levels of statistical significance. In historical Finland, the cohort sex ratio gauges selection against males in utero and predicts male infant mortality. The reproductive success findings, however, provide weak support for an evolutionarily adaptive explanation of male culling in utero.