Maternal investment in eggs is affected by male feet colour and breeding conditions in the blue-footed booby, <Emphasis Type="Italic">Sula nebouxii</Emphasis>

Females are expected to vary investment in offspring according to variables that may influence the offspring fitness in a way that optimises her inclusive fitness for a particular context. Thus, when sexual ornaments signal the quality of the male, females might invest in reproduction as a function of the attractiveness of their mate. We tested whether breeding conditions and male feet colour influence reproductive decisions of blue-footed booby females. In the blue-footed booby, male feet colour is a dynamic condition-dependent sexually selected trait that is related to paternal effort. During two consecutive years, an El Niño year (poor breeding conditions) and a year with good breeding conditions, we experimentally reduced male attractiveness by modifying their feet colour after the first egg was laid and recorded female investment in the second egg. We found that, relative to the first egg in the clutch, females laid heavier second eggs during the poor year than during the good year. Females paired with males with duller feet colour reduced second-egg mass and volume and delayed the laying of the second egg, independently of the year. Absolute yolk androstenedione (A4) concentration (but not testosterone, T) in second eggs was higher during a poor year than during a good year. Only during a year with poor breeding conditions, females paired with experimental males decreased the relative A4 concentration (but not T) in the second egg compared to control females. Thus, blue-footed booby females probably favour brood reduction by decreasing egg quality and increasing size asymmetry between chicks when the breeding and the mate conditions are poor.     

Sex allocation in the solitary bee Osmia cornuta: do females behave in agreement with Fisher's theory?

Fisher's theory of sex allocation predicts that, in a panmictic population, parental investment will be equally distributed between male and female progeny. Most studies on parental investment in nesting solitary bees and wasps use offspring or provision weight as estimators of parental investment and do not corroborate Fisher's theory. The measurement of parental investment may be confounded by several factors. First, the use of offspring or provision size does not account for seasonal variation in foraging costs associated with aging of nesting females. Second, provision or offspring size do not reflect parental investment associated with nest construction. In this two-year study we measured parental investment in a solitary bee. We calculated sex allocation using both provision weight and foraging time as parental investment estimators. Investment in pollen-nectar provisions decreased, while investment in mud structures (nest construction) increased, as the nesting period progressed. Overall investment in provisions per nest was ∼25 times higher than investment in mud. Pollen-nectar foraging trips became longer as the season progressed, but mud trip duration did not vary. Due to weather differences between years, more offspring per female were produced in the first year, but progeny sex ratio and mean offspring size of both sexes were similar between years. Mortality did not differ between sexes. As predicted by Fisher's theory, production cost ratios did not differ from 1 in either year, irrespective of the currency used to estimate parental investment (provision weight or foraging time). Our results strongly support Fisher's theory.     

The effect of a natural environmental disturbance on maternal investment and pup behavior in the California sea lion

Summary Observed changes in maternal investment due to an environmentally induced decrease in food supply (the 1983 El Ni?o-Southern Oscillation) are compared witha priori predictions for the California sea lion (Zalophus californianus). Changes in behavior, growth and mortality of off-spring were also examined. Data collected in the first two months postpartum for the years before (PRE), during (EN), and the two years after (POST1 and POST2) the 1983 El Ni?o indicate that females initiated postpartum feeding trips earlier during the food shortage, and spent more time away on individual feeding trips in both the El Ni?o year and the year after. Perinatal sex ratios (♀:♂) in the years PRE, EN, POST1 and POST2 were 1:1, 1.4:1, 1.1:1 and 1:1.4, respectively. Fewer copulations were observed during the El Ni?o year, but this difference was not statistically significant. Pups spent less time suckling in the food shortage year and the year following, but attempted to sneak suckle more. Pups were less active and played on land less in the El Ni?o and following year. Finally, maternal investment as measured by milk intake of offspring was decreased, pups grew more slowly, and suffered increased mortality during the food shortage year. Despite expected sex differences in maternal investment and pup behavior in response to food shortage, there were no sex-biased differences in response in either females or pups. As expected, the food shortage did not affect adult males since they migrate north during the non-breeding season where the environmental perturbation was less severe.     

Human preferences for facial masculinity change with relationship type and environmental harshness

In humans (Homo sapiens), sexual dimorphism in face shape has been proposed to be linked to quality in both men and women. Although preferences for high-quality mates might be expected, previous work has suggested that high quality may be associated with decreased investment in partnerships. In line with a trade-off between partner quality and investment, human females have been found to prefer higher levels of masculinity when judging under conditions where the benefits of quality would be maximised and the costs of low investment would be minimised. In this study, we examined facultative preferences for masculinity/femininity under hypothetical high and low environmental harshness in terms of resource availability in which participants were asked to imagine themselves in harsh/safe environments. We demonstrate that environmental harshness influences preferences for sexual dimorphism differently according to whether the relationship is likely to be short or long term. Women prefer less-masculine male faces and men prefer less-feminine female faces for long-term than short-term relationships under conditions of environmental harshness. Such findings are consistent with the idea that high-quality partners may be low investors and suggest that under harsh ecological conditions, both men and women favour a low-quality/high-investment partner for long-term relationships. For short-term relationships, where investment is not an important variable, preferences for sexual dimorphism were similar for the low and high environmental harshness conditions. These results provide experimental evidence that human preferences may be contingent on the environment an individual finds itself inhabiting.     

Maternal investment and sex-allocation in the Galapagos fur seal,Arctocephalus galapagoensis

Summary Maternal investment and sex-allocation were measured in a large, sexually dimorphic mammal, the Galapagos fur seal (Arctocephalus galapagoensis). The sex ratio at birth was 1.06. Males were always heavier than females and, at least initially, grew faster. Growth was variable from year to year suggesting energetic constraints on maternal investment. Sucking time conrrelated with milk intake. Mothers suckled yearling and 2-year-old sons more than daughters of the same age. Age at weaning appeared to be the same in both sexes or even slightly greater in males. No sex differences was found in mortality prior to weaning or in post-weaning dispersal. Birth rates of females with yearlings or 2-year-olds were significantly lower than those of females with no dependent young. Mothers invested more in sons than in daughters until weaning. It is unlikely that higher post-weaning investment in daughters balances the higher pre-weaning investment in sons. Data on sex ratio at birth, different growth rates, and weaning age of the sexes are typical of otariid seals as a group. The results of this study fit Maynard Smith's (1980) model of the evolution of sex allocation better than Fisher's (1930).     

Who cares? Individual contributions to pup care by breeders vs non-breeders in the cooperatively breeding banded mongoose (<Emphasis Type="Italic">Mungos mungo</Emphasis>)

Knowledge of the investment rules adopted by breeders and non-breeders, and the factors that affect them, is essential to understanding cooperative breeding as part of a life-history tactic. Although the factors that affect relative contributions to care of young have been studied in some cooperative bird species, there is little data on mammals, making coherent generalisations within mammals and across taxa difficult. In this study, we investigate individual contributions to pup escorting, a strong predictor of offspring provisioning, in the banded mongoose (Mungos mungo), a cooperatively breeding mammal in which reproductive skew is low. Contributions by those under a year old (which virtually never breed) increased with age and body weight but were generally low. Among older age classes (yearlings and adults), individuals that had not bred in the current litter generally contributed less to escorting than those that had bred (with the exception of yearling males). In addition, females that did not breed reduced their investment if they were heavy presumably because such females are more likely to breed in the following event and benefit from saving resources for this. The generally greater contributions by breeders in banded mongooses contrast with the recent findings in meerkats (Suricata suricatta), another obligatorily cooperative mongoose with similar group size but wherein reproductive skew is high. Our results suggest that relative contributions by breeders vs non-breeders are not dependent on group size but on the ratio of breeders to carers and the probability that non-breeders will breed in the near future.     

Maternal investment in rhesus macaques (Macaca mulatta): reproductive costs and consequences of raising sons

Maternal investment in offspring is expected to vary according to offspring sex when the reproductive success of the progeny is a function of differential levels of parental expenditure. We conducted a longitudinal investigation of rhesus macaques to determine whether variation in male progeny production, measured with both DNA fingerprinting and short tandem repeat marker typing, could be traced back to patterns of maternal investment. Males weigh significantly more than females at birth, despite an absence of sex differences in gestation length. Size dimorphism increases during infancy, with maternal rank associated with son’s, but not daughter’s, weight at the end of the period of maternal investment. Son’s, but not daughter’s, weight at 1 year of age is significantly correlated with adult weight, and male, but not female, weight accounts for a portion of the variance in reproductive success. Variance in annual offspring output was three- to fourfold higher in males than in females. We suggest that energetic costs of rearing sons could be buffered by fetal delivery of testosterone to the mother, which is aromatized to estrogen and fosters fat accumulation during gestation. We conclude that maternal investment is only slightly greater in sons than in daughters, with mothers endowing sons with extra resources because son, but not daughter, mass has ramifications for offspring sirehood. However, male reproductive tactics supersede maternal investment patterns as fundamental regulators of male fitness. Received: 23 July 1999 / Received in revised form: 23 February 2000 / Accepted: 13 March 2000     

Aging and improving reproductive success in horses: declining residual reproductive value or just older and wiser?

In many mammalian species, female success in raising offspring improves as they age. The residual reproductive value hypothesis predicts that each individual offspring will be more valuable to the mother as she ages because there is less conflict between the current and potential future offspring. Therefore, as mothers age, their investment into individual offspring should increase. Empirical evidence for an influence of declining residual reproductive value on maternal investment is unconvincing. Older mothers may not invest more, but may be more successful due to greater experience, allowing them to target their investment more appropriately (targeted reproductive effort hypothesis). Most studies do not preclude either hypothesis. Mare age significantly influenced maternal investment in feral horses living on the North Island of New Zealand. Older mares, that were more successful at raising foals, were more protective for the first 20 days of life, but less diligent thereafter. Total maternal input by older mothers did not seem to be any greater, but was better targeted at the most critical period for foal survival and a similar pattern was observed in mares that had lost a foal in the previous year. In addition, older mothers were more likely to foal in consecutive years, supporting the hypothesis that they are investing less than younger mares in individual offspring. Therefore, older mothers seem to become more successful by targeting their investment better due to experience, not by investing more in their offspring. Received: 22 September 1999 / Received in revised form: 30 October 1999 / Accepted: 6 November 1999     

Carbon Payments and Low‐Cost Conservation

Abstract: A price on carbon is expected to generate demand for carbon offset schemes. This demand could drive investment in tree‐based monocultures that provide higher carbon yields than diverse plantings of native tree and shrub species, which sequester less carbon but provide greater variation in vegetation structure and composition. Economic instruments such as species conservation banking, the creation and trading of credits that represent biological‐diversity values on private land, could close the financial gap between monocultures and more diverse plantings by providing payments to individuals who plant diverse species in locations that contribute to conservation and restoration goals. We studied a highly modified agricultural system in southern Australia that is typical of many temperate agriculture zones globally (i.e., has a high proportion of endangered species, high levels of habitat fragmentation, and presence of non‐native species). We quantified the economic returns from agriculture and from carbon plantings (monoculture and mixed tree and shrubs) under six carbon‐price scenarios. We also identified high‐priority locations for restoration of cleared landscapes with mixed tree and shrub carbon plantings. Depending on the price of carbon, direct annual payments to landowners of AU$7/ha/year to $125/ha/year (US$6–120/ha/year) may be sufficient to augment economic returns from a carbon market and encourage tree plantings that contribute more to the restoration of natural systems and endangered species habitats than monocultures. Thus, areas of high priority for conservation and restoration may be restored relatively cheaply in the presence of a carbon market. Overall, however, less carbon is sequestered by mixed native tree and shrub plantings.     

Intraspecific parasitism and split sex ratios in a monogynous and monandrous ant (Leptothorax nylanderi)

Sex ratios were bimodally distributed in a population of the monogynous and monandrous ant Leptothorax nylanderi during each of 3 study years. The population-wide investment ratios suggested worker control of sex allocation. Nest-level variation in the proportional investment in virgin queens was not affected by the presence or absence of a queen and only slightly by collecting year, but was correlated with nest size, total sexual investment and, unexpectedly, with differences in nestmate relatedness: small, low-investment nests and nests with several worker lineages produced male-biased sex ratios. Colonies containing several worker lineages arise from usurpation of mature colonies by unrelated founding queens and the fusion of unrelated colonies under strong nest site limitation. In contrast to facultatively polygynous and polyandrous species of social insects, where workers can maximize their inclusive fitness by adjusting sex ratios according to the degree of relatedness asymmetry, workers in mixed colonies of L. nylanderi do not benefit from manipulating sex allocation, as here relatedness asymmetries appear to be the same as in homogeneous colonies. Received: 7 December 1999 / Received in revised form: 29 February 2000 / Accepted: 13 March 2000     

Sex ratio and maternal rank in wild spider monkeys: when daughters disperse

Summary Data from a long-term field study of the spider monkey, Ateles paniscus, in Peru indicate that a strongly female-biased sex ratio exists from birth in this population. Of 46 infants born between July 1981 and June 1986, 12 were male, 32 were female and 2 were of undetermined sex. This effect is consistent between years as well, with more females than males born in each year of the study (Table 1). This bias is driven by the fact that low-ranking females produce daughters almost exclusively, while high-ranking females bias their investment somewhat less strongly towards sons (Table 2). The unusual pattern of female-biased maternal investment observed in this population of Ateles probably occurs for a combination of the following reasons: (1) maternal investment in individual male offspring is somewhat greater than in individual female offspring; (2) males remain with their natal groups, and the sons of high-ranking females are likely to be competitively superior to the sons of low-ranking females; (3) males compete for mates, and only the one or two most dominant males within a community are likely to achieve significant reproductive success. Two possible mechanisms of sex-ratio adjustment and the evidence for each are discussed.     

Inheritance of nest sites in female Columbian ground squirrels

Summary Home ranges, areas of intensive use, and locations of nest sites were determined for female Columbian ground squirrels and their daughters over seven years from a low-density population in southwestern Alberta. Adult females usually retained similar home ranges from year to year. However, they moved their nest sites more often than expected in years when a yearling daughter was present. Core areas and centres of activity also changed when the nest site was moved. Most daughters settled on their natal areas. The relinquishing of nest sites by females to their daughters is seen as a form of parental investment.     

Risk aversion and CO2 regulatory uncertainty in power generation investment: Policy and modeling implications

We consider a simulation of risk-averse producers when making investment decisions in a competitive energy market, who face uncertainty about future regulation of carbon dioxide emissions. Investments are made under regulatory uncertainty; then the regulatory state is revealed and producers realize returns. We consider anticipated taxes, grandfathered permits and auctioned permits and show that some anticipated policies increase investment in the relatively dirty technology. Beliefs about the policy instrument that will be used to price carbon may be as important as certainty that carbon will be priced. More generally, a failure to consider risk aversion may bias policy analysis for the power sector.     

Relationship between body size, provisioning rate, longevity and reproductive success in females of the solitary bee Osmia cornuta

Body size has often been related to reproductive success in bees and wasps. The objective of this 3-year study was to analyze the relationship between nesting female body size, provisioning rate and longevity and their effect on several traits related to parental investment and reproductive success in the solitary bee Osmia cornuta. Body size was not correlated to longevity, and it was only correlated to provisioning rate in the third year (with poor weather conditions during nesting). Variation in fecundity, offspring size and offspring mortality was not well explained by nesting female body size in any of the 3 years. However, in the third year, small females biased their investment toward males, the sex requiring smaller pollen–nectar provisions. Large females were more successful usurpers of other females' nests, but fecundity of usurpers was no higher than fecundity of nonusurpers. Large females were more likely to establish at the release site, probably in relation to size-dependent vigor at emergence. A review of the literature on parental investment in solitary aculeate Hymenoptera showed a stronger relationship between body size and reproductive success in wasps than in bees. In O. cornuta, fecundity was strongly related to longevity and provisioning rate in all 3 years. Offspring size was associated with provisioning rate in 1 year, when females with higher provisioning rates tended to produce larger sons and daughters. Both longevity and provisioning rate appeared to be strongly conditioned by stochastic events.     

The relationship between maternal ornamentation and feeding rate is explained by intrinsic nestling quality

In altricial birds, parental feeding is essential, and its amount may depend on the quality of both parents. A relationship between parental quality and feeding rate is generally attributed to an active adjustment by parents in order to retain good quality mates or ensure high fitness through raising high-quality offspring. However, the behaviour and need of young may also change with parental quality, and this may affect parental behaviour. A further problem is that most studies have investigated post-hatching parental investment in relation to the secondary sexual signals of males, but not females. In a cross-fostering experiment, we examined the feeding rates of rearing parents in relation to the size and ornamentation of both original and rearing parents in collared flycatchers (Ficedula albicollis). Using this setup, we could examine whether the observed feeding patterns were the results of the decision of the parents based on their own and their partner’s traits or the constraints imposed by the behaviour or need of offspring. When correcting for clutch size and year, we found that feeding rate of both foster parents correlated with the wing patch size of the original female. This implies that original maternal quality had an offspring-mediated indirect effect on investment of foster parents, that is intrinsic nestling quality may constrain parental feeding decisions. This explanation should not be overlooked in future studies on preferential parental investment, and our results also point out that maternal ornaments deserve more attention in such studies.     

Parental investment without kin recognition: Simple conditional rules for parent-offspring behavior

Species differ widely with regard to parental investment strategies and mechanisms underlying those strategies. The passing of benefits to likely offspring can be instantiated with a number of different computational and behavioral systems. We report results from an agent-based model in which offspring maintain proximity with parents and parents transmit benefits to offspring without the capacity of either parent or offspring to 'recognize' one another. Instead, parents follow a simple rule to emit benefits after reproducing and offspring follow a simple rule of moving in the direction of positive benefit gradients. This model differs from previous models of spatial kin-based altruism in that individuals are modeled as having different behavioral rules at different life stages and benefits are transmitted unidirectionally from parents to offspring. High rates of correctly directed parental investment occur when mobility and sociality are low and parental investment occurs over a short period of time. We suggest that strategies based on recognition and bonding/attachment might serve to increase rates of correctly directed parental investment under parameters that are shown here to otherwise lead to high rates of misdirected and wasted parental investment.     

Male-biased maternal expenditure and associated costs in fallow deer

This study tested whether fallow deer mothers, Dama dama, bias their investment towards sons and, thus, whether sons are more costly to produce than daughters. Young (2 years) and old (≥3 years) hinds were analysed separately. Old hinds who raised sons accumulated less body mass than those who raised daughters, during the period between late gestation and the end of lactation. This difference in body mass persisted to the following spring. Mothers who had raised sons gave birth later and their offspring's pre-winter mass was lower the following year than for mothers who had raised daughters. These results indicate higher expenditure for hinds who raise sons and support theories of male-biased maternal investment. However, young mothers with sons and those with daughters did not differ in reproductive performance the following year. One reason might be that young mothers are close to the maximum level of maternal expenditure, since they are still growing, and cannot invest any extra resources in sons. Received: 28 August 1997 / Accepted after revision: 5 April 1998     

Exploring price effects on the residential water conservation technology diffusion process: a case study of Tianjin city

Reforms of the water pricing management system and the establishment of a flexible water pricing system are significant for cities in northern China to tackle their critical water issues. The WATAP (Water conservation Technology Adoption Processes) model is developed in order to capture the water conservation technology adoption process under different price scenarios with disaggregate water demands down to the end use level. This model is explicitly characterized by the technological selection process under maximum marginal benefit assumption by different categories of households. In particular, when households need to purchase water devices in the provision market with the consideration of complex factors such as the life span, investment and operating costs of the device, as well as the regulated water price by the government. Applied to Tianjin city, four scenarios of water price evolutions for a long-term perspective (from year 2011 to 2030) are considered, including BAU (Business As Usual), SP1 (Scenario of Price increase with constant annual rate), SP2 (Scenario of Price increase every four years) and SP3 (Scenario of Price increase with affordable constraint), considering many factors such as historic trends, affordability and incentives for conservation. Results show that on aggregate 2.3%, 11.0% and 18.2% of fresh water can be saved in the residential sector in scenario SP1, SP2 and SP3, respectively, compared with the BAU scenario in the year 2030. The water price signals can change the market shares of different water appliances, as well as the water end use structure of households, and ultimately improve water use efficiency. TheWATAP model may potentially be a helpful tool to provide insights for policy makers on water conservation technology policy analysis and assessment.     

Parental investment without kin recognition: simple conditional rules for parent–offspring behavior

Species differ widely with regard to parental investment strategies and mechanisms underlying those strategies. The passing of benefits to likely genetic offspring can be mediated through a number of different computational and behavioral systems. We report results from an agent-based model in which offspring maintain proximity with parents and parents transmit benefits to offspring without the capacity of either parent or offspring to “recognize” one another. Instead, parents follow a simple rule to emit benefits after reproducing and offspring follow a simple rule of moving in the direction of positive benefit gradients. This model differs from previous models of spatial kin-based altruism in that individuals are modeled as having different behavioral rules at different life stages and benefits are transmitted unidirectionally from parents to offspring. High rates of correctly directed parental investment occur when mobility and sociality are low and parental investment occurs over a short period of time. We suggest that strategies based on recognition and bonding/attachment might serve to increase rates of correctly directed parental investment under parameters that are shown here to otherwise lead to high rates of misdirected and wasted parental investment.     

Optimal maternal investment in the dung beetle<Emphasis Type="Italic"> Onthophagus taurus</Emphasis>?

Optimal investment theory is based on the assumption that the proximate constraint acting on parental investment is resource based. A trade-off between per offspring investment and total investment seems intuitive. Consequently, a parents investment strategy is expected to represent a trade-off between the benefits of investment for current offspring and the costs to future reproduction for parents. In this study, we provide clear evidence that the costs and benefits of maternal provisioning in the dung beetle Onthophagus taurus influence the amount of provisions provided by the mother. Horse dung is typically of a higher nutritional value than cow dung and females were shown to provide 20% less dung to offspring when provisioning with horse dung. By reducing their investment per offspring and exhibiting a clear preference to provision offspring with horse dung, females were able to produce significantly more offspring. Females provisioning with horse dung received greater fitness returns per unit of investment and experienced lower provisioning costs, in terms of the minimum amount of dung required to produce a surviving offspring, than females provisioning with cow dung. Females provisioning in soil of low moisture content were found to have higher tunneling costs than those provisioning in soil of high moisture content, while the fitness returns per unit of investment did not differ. We adopted a marginal value theorem (MVT) approach to calculate the theoretical optimal level of investment for each dung type and for each soil moisture. Predicted levels of provisioning were lower for horse dung than for cow dung and for moist soil than for dry soil. Therefore, the results of this study are in qualitative agreement with MVT predictions and provide empirical support for the proposal that females can adaptively adjust their level of investment in response to resource and/or habitat quality. However, the theoretically predicted optimal investment yielded a poor quantitative fit with our observed levels of investment, with females providing over twice the investment predicted by the MVT approach. We suggest that this difference may reflect either our inability in directly quantifying all the necessary costs and benefits of investment in O. taurus and/or the applicability of the underlying assumptions of MVT.Communicated by D. Gwynne