Lactation,maternal behavior and infant growth in common marmoset monkeys (<Emphasis Type="Italic">Callithrix jacchus</Emphasis>): effects of maternal size and litter size

We examined the relationship between lactation performance and infant growth in a captive population of common marmosets (Callithrix jacchus) that varied in both maternal and litter size. Though common marmosets display a typical primate pattern of dilute milk and relatively slow infant growth rates (factors associated with low daily lactation investment and minimal maternal size effects), we hypothesized that the marmoset's small body size would make lactation investment more sensitive to maternal size than is true for larger-bodied primates. Smaller mothers rearing twins had lower milk fat, lower gross energy in milk samples collected in mid to late lactation and lower nursing-bout frequencies than did large mothers nursing twins. Lactation performance and maternal behavior did not differ between large and small mothers when rearing singletons, with a single exception: small mothers had a lower gross energy in mid-lactation milk samples. Relative growth rates in twins but not singletons were affected by maternal size, such that small mothers supported more growth per infant when rearing singletons while large mothers supported more growth per infant when rearing twins. Among the larger mothers, only, older mothers supported somewhat, though not significantly, less growth per infant, regardless of litter size. Twin infants of small mothers appeared to respond to below-optimal levels of milk yield by initiating maternal carrying less often. The relative energy intake of mothers was similar regardless of litter or maternal size. Small mothers rearing twins drew more heavily on reserves, reflected in a linear weight loss during lactation; however, the reserves drawn upon were inadequate to meet the lactation demand, resulting in lower milk energy output. In addition, small mothers rearing twins were more likely to be ill and less likely to be fertile in the year following lactation than were large mothers of twins or mothers of any size rearing singletons.     

Costs and benefits of communal rearing to female prairie voles (<Emphasis Type="Italic">Microtus ochrogaster</Emphasis>)

Communal rearing of offspring may help mothers maximize their investment in offspring at a reduced cost to their own bodily condition, thus maximizing their potential for reproductive success. The objective of this study was to quantify the costs and benefits of communal rearing to prairie vole (Microtus ochrogaster) pups and mothers. Mothers were assigned to one of three social units: solitary mothers, singularly breeding groups (i.e. one mother and one non-reproductive sister) and plurally breeding groups (i.e. two lactating sisters). For each type of social unit, some replicates were provided with food ad libitum, while others were provided with limited food. The body mass of focal mothers (i.e. the first mother to produce a litter) was a significant predictor of pup growth. Regardless of food availability, litters of focal mothers in plurally breeding groups gained more weight than litters reared by solitary mothers. Pups reared in singularly breeding groups were intermediate in weight gain, but did not gain significantly more weight than solitary offspring. There was no difference in the body mass of focal mothers from each type of social unit, regardless of food availability. Within plurally breeding groups, the weight gain of the two litters and body mass of focal and second mothers did not differ. However, focal mothers from plurally breeding groups nursed fewer pups than solitary mothers and also fewer pups than their nestmates when food was limited. Our results suggest that plural breeding results in greater fitness to mothers than solitary and singular breeding.Communicated by E. Korpimäki     

Lactation performance of guinea pigs (Cavia porcellus) does not respond to experimental manipulation of pup demands

To ensure proper development of young, mothers should react to offspring signals of need. Studies of such parent-offspring interaction often manipulated litter size to measure effects of changed offspring food demand. We used the extreme precociality of guinea pig (Cavia porcellus) pups to increase offspring demand without changing any other litter characteristic. As pups contribute to their own energy demands from birth by independent feeding, their food demand can be increased by withholding solid food early in lactation. We studied whether mothers reacted to increased food demand of offspring by enhanced parental care, especially by changes in the pattern of milk production and nursing. Pups deprived of solid food early in lactation grew more slowly and were in poorer body condition than pups in control litters, even after the former had access to solid food in late lactation. Mothers of deprived young reacted to offspring long-term need by maintaining nursing behaviour for longer than control mothers. However, this change in behaviour did not occur early in lactation when pup short-term need was greatest nor did it result in increased milk transfer at any time. Energy allocation of mothers measured by changes in their food intake, maternal body reserves, and milk production stayed the same whether offspring had early access to solid food or not. Thus mothers did not increase energy allocation to pups even though they apparently had information about the pups' poor state.     

Early maternal investment in male and female African elephant calves

Summary The suckling behaviour of 130 freeranging elephant calves aged between birth and 4.5 years old was examined in Amboseli National Park, Kenya. Analyses of frequencies of suckling and durations of suckling bouts showed that males attempted to suckle more often, were more successful at their attempts, and as a result were estimated to have a higher milk intake than did female calves. Mothers were equally tolerant of their sons' and daughters' demands to suckle at young ages, but were less tolerant of their older sons' demands. The growth rates of males based on hind footprint length were faster than those of females from birth onwards. During drought years with low food availability, male calf survivorship in the first year was lower than that of female calves. During wet years, there was little difference between sexes in survivorship. It appeared that during dry years mothers were unable to sustain milk production at a level that met the metabolic requirements of their sons, and as result male calves were more likely to die. Females with a surviving son tended to have a longer interbirth interval than did females with a surviving daughter. We suggest that greater early maternal investment in male calves occurs because, in the highly-competitive polygynous mating system of elephants, size in adult male elephants is an important factor in mating success.     

Harem choice and breeding experience of female southern elephant seals influence offspring survival

Female mammals can increase their lifetime fitness through modification of investment potential and by providing better rearing environments with improved breeding experience. We examined the relationships between reproductive fitness and the behavioural decisions that female southern elephant seals (Mirounga leonina) made during the breeding season. We examined whether mother age and breeding experience influenced reproductive success (measured as 1st-year survival probability), and whether there was a change in the choice of harem size with increasing age. Pups produced by young mothers had lower 1st-year survival probability than pups produced by older mothers. A significant increase in mean female mass with age required an analysis of both these effects on offspring survival. There was a significant positive effect of both female age and mass, and the interaction between the two, on 1st-year pup survival. The proportion of young mothers (<5 years old) decreased and the proportion of older mothers (>6 years old) increased with increasing harem size (harems surveyed from 1997 to 2001). Females chose larger harems in which to breed as they aged. Females demonstrated fidelity to breeding areas among successive breeding seasons, with older females displaying greater breeding-site fidelity than younger females. The mean number of previous breeding attempts per female within a harem (breeding experience) increased significantly with increasing harem size. Breeding females returned to breed later in the breeding season as they aged—we hypothesize that young, subordinate females gain a priority advantage by returning earlier. These results lend support to the hypothesis that there are fitness advantages, in terms of offspring survival, that are conferred to females that breed in successively larger harems with age. Potential mechanisms that select for females to improve their breeding conditions include improved mate selection and the avoidance of conspecific harassment in harems.Communicated by F. Trillmich     

Allocation in reproduction is not tailored to the probable number of matings in common toad (<Emphasis Type="Italic">Bufo bufo</Emphasis>) males

The theory of life history evolution assumes trade-offs between competing fitness traits such as reproduction, somatic growth, and maintenance. One prediction of this theory is that if large individuals have a higher reproductive success, small/young individuals should invest less in reproduction and allocate more resources in growth than large/old individuals. We tested this prediction using the common toad (Bufo bufo), a species where mating success of males is positively related to their body size. We measured testes mass, soma mass, and sperm stock size in males of varying sizes that were either (1) re-hibernated at the start of the breeding season, (2) kept without females throughout the breeding season, or (3) repeatedly provided with gravid females. In the latter group, we also estimated fertilization success and readiness to re-mate. Contrary to our predictions, the relationship between testes mass and soma mass was isometric, sperm stock size relative to testes mass was unrelated to male size, fertilization success was not higher in matings with larger males, and smaller males were not less likely to engage in repeated matings than larger males. These results consistently suggest that smaller males did not invest less in reproduction to be able to allocate more in growth than larger males. Causes for this unexpected result may include relatively low year-to-year survival, unpredictable between-year variation in the strength of sexual selection and low return rates of lowered reproductive investment.     

Factors influencing length of maternal care in brown bears (<Emphasis Type="Italic">Ursus arctos</Emphasis>) and its effect on offspring

Length of maternal care, i.e. the interval between successfully raised litters, is the most important factor explaining the variation in reproductive rate among brown-bear (Ursus arctos) populations. In this paper, we examine the variation in length of maternal care in radio-marked brown bears and its effect on their offspring in northern Sweden. Young stayed with their mothers for 1.4–1.5 or 2.4–2.5 (in one case 3.5) years and were weaned with body masses varying from 17 to 69 kg. The probability of yearling litters staying with their mother for a 2nd year increased with decreasing yearling body mass, and was higher for litters with two offspring than for litters with one or three to four offspring. Staying with their mothers for a 2nd year had a positive effect on mass gain in yearlings and this effect was more pronounced in litters with two than three to four offspring. Body mass of 2-year-olds was not related to age of weaning, suggesting that keeping offspring for an additional year mainly compensated for low yearling body mass. If large offspring body mass positively affects later offspring survival and reproduction, mothers may be able to optimize the length of maternal care according to the litter size and the size of their yearlings.Communicated by F. Trillmich     

Sibling competition in guinea pigs (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): scrambling for mother’s teats is stressful

Lactation is the most energy-intense period in the life of a female mammal. This can cause severe conflict between mother and offspring over the duration of lactation but also between siblings over the amount of milk each pup gets from its mother. Thus, competitive interactions between siblings are expected, and competition is likely to increase with litter size, particularly in species where the number of offspring exceeds the number of teats. We studied sibling competition in the domestic guinea pig (Cavia aperea f. porcellus), which has two teats, but frequently bears litters of up to five pups. By cross-fostering we created non-competition (control) litters with two pups and competition litters with four pups and observed nursing behaviour on days 5, 10, 15 and 20 postpartum. Pups of larger litters had lower growth rates, indicating increased competition among siblings in these litters. Pups of larger litters had to wait longer for access to a teat and spent less time suckling than pups of smaller litters but ate more solid food instead. Additionally, we manipulated the individual short-term need of pups by separating half of the pups of each litter for 2 h from their mothers before observation. Within a litter, hungry pups achieved access to milk faster and spent more time suckling than non-hungry pups. Pups competed mostly by scramble competition. Aggressive interactions occurred only in large litters. Pups of large litters had higher cortisol levels than pups in small litters. These effects decreased with age as pups became increasingly independent of maternal milk. Pup behaviour appears to fit better with models of scramble competition than with those of honest signalling. This contribution is part of the special issue “Sibling competition and cooperation in mammals” (guest editors: Robyn Hudson and Fritz Trillmich).     

The demographic consequences of the cost of reproduction in ungulates

The cost of reproduction can generate covariation between demographic rates that can potentially influence demography and population dynamics in long-lived iteroparous species. However, there has been relatively little work linking the survival cost of reproduction and population dynamics. The apparent scarcity of information on this important link is potentially due to covariation between vital rates, which can substantially influence fluctuations in population size. In this paper we examine the opportunity for survival costs of reproduction to leave a dynamic signature using a simulation model based broadly on an ungulate life history. We find that an increase in the cost delays the onset of reproduction and reduces reproductive rates of young, but not of prime-age, females. Accordingly, the number of offspring produced declines and the interval between reproductive events increases among young females experiencing high cost. These effects are translated to an age structure skewed toward young ages and reduced population density. These results suggest that, by delaying reproduction when conditions deteriorate, females protect their survival during the critical first three years of life, after which the negative effect of reproduction on survival declines. Unless conditions for reproduction are severe, it is not profitable to delay reproduction beyond age 3 years due to the high risk of death before having a chance to reproduce. We also demonstrate that lack of adjustment of reproductive strategies to elevated levels of the cost of reproduction, for example due to rapid changes in environmental conditions, results in lower average density and longevity compared to females that have sufficient time to adjust to changes in the cost. This suggests that even moderate costs of reproduction may have a major negative effect on population dynamics of ungulates.     

Variation in litter size: a test of hypotheses in Richardson's ground squirrels

We studied litter size variation in a population of Richardson's ground squirrels (Spermophilus richardsonii) in Alberta, Canada, from 1987 to 2004. Litter size at first emergence of juveniles from the natal burrow ranged from 1 to 14; the most common litter sizes, collectively accounting for 41.0% of 999 litters, were 6 and 7. The number of offspring surviving to adulthood (attained on emergence from hibernation as yearlings) increased with increasing litter size, a result that was not predicted by Lack's "optimal litter size" hypothesis, Mountford's "cliff-edge" effect, or the "bad-years" effect. Contrary to the negative effects predicted by the "cost of reproduction" hypothesis, litter size had no significant influence on survival of mothers to the subsequent year or on the size of the subsequent litter. Rather, our results best fit the predictions of the "individual optimization" hypothesis, which suggests that litter size is determined by the body condition and environmental circumstances of each mother. Supporting this hypothesis, survival of individual offspring was not significantly associated with litter size. Additionally, year-to-year changes in maternal body mass at mating were positively associated with concurrent changes in litter size (r = 0.56), suggesting that litter size depends on the body condition of the mother. Because the mean number of recruits to adulthood increased as litter size increased (r2 = 0.96) and litter size increased with maternal condition, offspring productivity was greater for mothers in better body condition.     

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.     

Weaning in the guinea pig (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): Who decides and by what measure?

Offspring should be selected to influence maternal effort to maximize their own fitness, whereas mothers are selected to limit investment in present progeny. In mammals, this leads to a conflict over the amount of milk provided and the timing of weaning. The intensity and time course of such conflict has so far mostly been investigated experimentally in altricial rodents. However, it is expected that offspring options for conflict will depend on developmental state. We therefore investigated in the highly precocial domestic guinea pig (Cavia aperea f. porcellus) who decides over nursing performance and weaning and how pup state influences these decisions. Specifically, we tested whether a threshold mass of pups predicts weaning time. By exchanging older litters against neonates and vice versa, we produced a situation in which females differed in lactational stage from the cross-fostered pups. Our results indicate that females decide about the timing of weaning, as cross-fostered younger pups were weaned at a much younger age than controls and older pups benefited from continuing lactation of foster mothers. Growth rates did not differ in the treatment groups, and different weaning ages resulted in differing weaning mass refuting the hypothesis that weaning is based on a threshold mass of offspring. This constitutes clear evidence that in a precocial rodent, the guinea pig, decisions about maternal care are primarily determined by maternal state and little influenced by pup state despite the extreme precociality of offspring. We suggest that precocial pups show little resistance to early weaning when food is abundant, as they reach sufficient nutritional independence by the middle of lactation to enable independent survival.     

The relationship between maternal phenotype and offspring quality: do older mothers really produce the best offspring?

Maternal effects are increasingly recognized as important drivers of population dynamics and determinants of evolutionary trajectories. Recently, there has been a proliferation of studies finding or citing a positive relationship between maternal size/age and offspring size or offspring quality. The relationship between maternal phenotype and offspring size is intriguing in that it is unclear why young mothers should produce offspring of inferior quality or fitness. Here we evaluate the underlying evolutionary pressures that may lead to a maternal size/age-offspring size correlation and consider the likelihood that such a correlation results in a positive relationship between the age or size of mothers and the fitness of their offspring. We find that, while there are a number of reasons why selection may favor the production of larger offspring by larger mothers, this change in size is more likely due to associated changes in the maternal phenotype that affect the offspring size-performance relationship. We did not find evidence that the offspring of older females should have intrinsically higher fitness. When we explored this issue theoretically, the only instance in which smaller mothers produce suboptimal offspring sizes is when a (largely unsupported) constraint on maximum offspring size is introduced into the model. It is clear that larger offspring fare better than smaller offspring when reared in the same environment, but this misses a critical point: different environments elicit selection for different optimal sizes of young. We suggest that caution should be exercised when interpreting the outcome of offspring-size experiments when offspring from different mothers are reared in a common environment, because this approach may remove the source of selection (e.g., reproducing in different context) that induced a shift in offspring size in the first place. It has been suggested that fish stocks should be managed to preserve these older age classes because larger mothers produce offspring with a greater chance of survival and subsequent recruitment. Overall, we suggest that, while there are clear and compelling reasons for preserving older females in exploited populations, there is little theoretical justification or evidence that older mothers produce offspring with higher per capita fitness than do younger mothers.     

Maternal quality and differences in milk production and composition for male and female Iberian red deer calves (Cervus elaphus hispanicus)

Several theories predict a sex-biased investment either through unbalanced sex ratios in offspring or through differences in provisioning. According to them, one would expect an optimisation in indirect fitness, or else a compensation for increased mortality of one sex. In addition, biases in provisioning may also arise as a consequence of weight-dependent non-adaptive nutrient demands by offspring. This study examines milk provisioning and sex biases in offspring sex ratio together with maternal quality variables. Mothers of higher quality (weight and age) showed greater milk provisioning ability (in terms of production) resulting in greater calf weight gain. Mothers of sons produced greater yields of milk, milk protein, fat and lactose than mothers of daughters, and increased percentage of protein after controlling for higher male birth weight. In contrast, mothers of males did not differ from mothers of females in age or any body weight variables related to maternal quality. These results suggest that differences in milk production and composition for sons and daughters are rather a mechanism to optimise indirect fitness than a mechanism to compensate for increased mortality in male calves, or a consequence of greater weight-dependent nutrient demands by heavier male calves. Results also suggest that biases in milk provisioning may occur without biases in offspring sex ratio, and furthermore, in contrast to the prediction that biases should be relative to the mean investment of the population, that milk provisioning biases might not be relative.Communicated by F. Trillmich     

Mixed sex allocation strategies in a polytocous mammal,the house mouse (<Emphasis Type="Italic">Mus musculus</Emphasis>)

The issue of adaptive adjustment of offspring sex ratio (proportion of male births) in polytocous mammals, producing several offspring per litter, is controversial because females of these species can maximize their fitness mainly by adjusting offspring number. To address this issue, we examined the effect of maternal condition at mating, experimentally decreased by pre-mating food restriction, on the sex ratio variation in 137 female mice. We tested two basic sex allocation hypotheses plausible for polytocous mammals: (1) the Myers hypothesis, predicting that cheaper sex should be favored in poor environmental conditions to maximize offspring number; and (2) the Williams hypothesis, predicting maximum fitness returns by adjusting size- and sex-specific composition of the litter according to the maternal condition. The food-restricted mothers produced larger litters with a higher proportion of cheaper daughters than the control mothers. By contrast, the control mothers optimized size and sex composition of the litter according to their weight at mating. In addition, the offspring of the food-restricted mothers suffered less from pre-weaning mortality than those of the control mothers. Therefore, when comparing the groups, the Myers hypothesis had a general significance while the Williams hypothesis was plausible only for the control mothers. Furthermore, some of the food-restricted mothers partly coped with the pre-mating food restriction and increased the proportion of sons in the litter with the increasing maternal weight loss (during the period of food restriction). The sex ratio variation was thus a result of three sex allocation strategies depending on the maternal condition at mating.     

A model for energetics and bioaccumulation in marine mammals with applications to the right whale.

We present a dynamic energy budget (DEB) model for marine mammals, coupled with a pharmacokinetic model of a lipophilic persistent toxicant. Inputs to the model are energy availability and lipid-normalized toxicant concentration in the environment. The model predicts individual growth, reproduction, bioaccumulation, and transfer of energy and toxicant from mothers to their young. We estimated all model parameters for the right whale; with these parameters, reduction in energy availability increases the age at first parturition, increases intervals between reproductive events, reduces the organisms' ability to buffer seasonal fluctuations, and increases its susceptibility to temporal shifts in the seasonal peak of energy availability. Reduction in energy intake increases bioaccumulation and the amount of toxicant transferred from mother to each offspring. With high energy availability, the toxicant load of offspring decreases with birth order. Contrary to expectations, this ordering may be reversed with lower energy availability. Although demonstrated with parameters for the right whale, these relationships between energy intake and energetics and pharmacokinetics of organisms are likely to be much more general. Results specific to right whales include energy assimilation estimates for the North Atlantic and southern right whale, influences of history of energy availability on reproduction, and a relationship between ages at first parturition and calving intervals. Our model provides a platform for further analyses of both individual and population responses of marine mammals to pollution, and to changes in energy availability, including those likely to arise through climate change.     

Current indirect fitness benefits associated with philopatry in juvenile prairie voles

Summary Prairie vole (Microtus ochrogaster) family groups were examined to determine possible indirect fitness benefits from the presence of juvenile helpers at the natal nest. The reproductive performance of family groups retaining two juveniles was compared with that of families in which all juveniles were removed when the subsequent litter was born. Litter sizes at birth of the second litter and pup survival rates were the same in the two treatments. Offspring quality was affected by the presence of juveniles however. Pups reared with juveniles weighed 13% more at weaning than pups reared without juveniles. Pups also opened their eyes sooner when juveniles were present. Differences in growth and development may have been affected by the amount of time pups were alone in the nest since pups in families with juveniles were left alone less frequently than were pups without juveniles. Maternal behavior patterns were not affected by the presence of juveniles. In contrast, fathers in families with juveniles spent more time in non-parental behaviors such as feeding, drinking, and foraging. In families with large litters, mothers delivered a subsequent litter sooner if juveniles were present. Subsequent litter sizes were the same in both treatments. Overall, both infants and parents benefited from the presence of juveniles, suggesting that helping may enhance the helper's indirect fitness in multiple ways.     

Experimental evidence for food limitation and sex-specific strategies of American kestrels (<Emphasis Type="Italic">Falco sparverius</Emphasis>) provisioning offspring

For birds with altricial young, the brood-rearing period is one of the most energetically expensive periods in their lives and may be the bottleneck for fitness. Because parents are expected to be prudent in allocating resources between reproduction and self-maintenance, food supply should be an important factor determining reproductive decisions during brood rearing. Parents with abundant food are expected to have enhanced fitness because they are able to reduce their work rates and increase their own survival chances, because their offspring may be of higher quality, or some combination of these. However, few studies have simultaneously documented all of these variables in a single investigation. We performed a food supplementation experiment to test how food supply influenced provisioning decisions by parent American kestrels (Falco sparverius). Female kestrels showed a strong response to extra food and reduced their provisioning rates. As a result, supplemented females had higher return rates than control females, suggesting significant effects of food on female survival. Because females used extra food to increase their fitness, our results suggest that kestrels raising offspring are limited by food. Male kestrels whose nests were supplemented also responded to extra food by reducing provisioning, but to a much lesser extent than their mates. Male parents did not appear to benefit from supplementation, as their return rates were similar to control males. The total amount of food received on a daily basis by nestlings was similar between supplemented and control nests. Supplemented offspring therefore did not fledge in better condition or have higher survival rates than control nestlings; the only significant factor consistently affecting offspring condition and survival was weather.     

Siblicide in Serengeti spotted hyenas: a long-term study of maternal input and cub survival

In the Serengeti National Park, Tanzania, large fluctuations of prey abundance alters the frequency at which spotted hyena (Crocuta crocuta) cubs are nursed and thus the total level of maternal input available to them. Maternal input is high when mothers feed on high densities of locally available migratory herbivores and low when mothers travel up to 70 km to forage. Using data from 19 cub cohorts on the incidence of siblicide (from monitoring the survival of 609 cubs in twin litters) and cub growth rates (from 195 cubs in twin litters) as a measure of maternal input, we demonstrate that the incidence of siblicide increased as average cohort growth rate declined. In total, there were 37 siblicides in 384 litters (9% of litters). When both cubs were alive, total maternal input in siblicidal litters was significantly lower than in non-siblicidal litters and the mean share of the dominant sib of 64.6% was significantly higher than the mean of 52.1% for dominant sibs of non-siblicidal litters. After siblicide, growth rates of siblicide victors substantially increased, demonstrating that mothers did not reduce maternal input after litter reduction. As a result, siblicide victors achieved a long-term growth rate similar to that of singletons and thus significantly increased their expected survival. We conclude that in spotted hyenas, high maternal input in lactation has favoured the evolution of facultative siblicide in populations inhabiting areas with low or fluctuating food resources.     

Age-related foetal sex ratio bias in Iberian red deer (<Emphasis Type="Italic">Cervus elaphus hispanicus</Emphasis>): are male calves too expensive for growing mothers?

Several theories predict offspring biases towards males or females with increasing reproductive resources of the mother to maximize reproductive returns by offspring, or as a result of prohibitive cost of the most expensive sex for young mothers or those in poor condition. This study examines foetus sex of 221 harvested hinds in a food-supplemented game estate for 10 years, according to hind age class (yearlings, subadults or adults), precise age, body mass and condition, and jaw length. A logistic model showed that hinds had a greater probability of bearing a male foetus with increasing age class, but not with any other variable. The greatest bias was found in yearling hinds. After controlling for age class and mass, jaw length was smaller in pregnant compared to non-pregnant yearlings and subadults, which suggests a trade-off between reproduction and growth. The bias towards females in yearlings increased as gestation proceeded, which suggests that the bias might be a result of selective abortion of male foetuses. Although results do not exclude an investment in males to increase number of grand-offspring, they suggest that young hinds may produce daughters as a trade-off between low energetic-cost offspring and their need to grow.Communicated by S. Krackow