Growth and ontogeny of sexual size dimorphism in the mandrill (Mandrillus sphinx)

We present body mass (N = 419) and crown-rump length (CRL, N = 210) measurements from 38 male and 49 female mandrills born into a semifree-ranging colony in order to describe growth from birth to adulthood, and to investigate maternal influences upon growth. Adult male mandrills are 3.4 times the body mass, and 1.3 times the CRL, of adult females. Body mass dimorphism arises from a combination of sex differences in length of the growth period (females attain adult body mass at 7 years, males at 10 years) and growth rate. Both sexes undergo a subadult growth spurt in body mass, and this is much more dramatic in males (peak velocity 551 g/months +/- 89 SEM at 84-96 months). CRL dimorphism arises from bimaturism (females attain adult CRL at 6 years, males after 10 years), and neither sex shows a particular subadult growth spurt in CRL. Sexual size dimorphism thus represents important time and metabolic costs to males, who mature physically approximately 3-4 years after females. Considerable interindividual variation occurs in the size-for-age of both sexes, which is related to maternal variables. Older mothers have heavier offspring than do younger mothers, and higher-ranking mothers have heavier offspring than do lower ranking mothers. Mass advantages conferred upon offspring during lactation by older and higher-ranking mothers tend to persist postweaning in both sexes. Thus maternal factors affect reproductive success in both sexes, influencing the age at which offspring mature and begin their reproductive career.     

Mothers determine offspring size in response to own juvenile growth conditions

Through non-genetic maternal effects, mothers can tailor offspring phenotype to the environment in which young will grow up. If juvenile and adult ecologies differ, the conditions mothers experienced as juveniles may better predict their offspring's environment than the adult environment of mothers. In this case maternal decisions about investment in offspring quality should already be determined during the juvenile phase of mothers. I tested this hypothesis by manipulating juvenile and adult maternal environments independently in a cichlid fish. Females raised in a poor environment produced larger young than females raised without food limitations, irrespective of the feeding conditions experienced during adulthood. This maternal boost was due to a higher investment in eggs and to faster larval growth. Apparently, mothers prepare their offspring for similar environmental conditions to those they encountered as juveniles. This explanation is supported by the distribution of these fishes under natural conditions. Juveniles live in a different and much narrower range of habitats than adults. Therefore, the habitat mothers experienced as juveniles will allow them to predict their offspring's environment better than the conditions in the adult home range.     

Persistent maternal effects on juvenile survival in North American red squirrels

Maternal effects can have lasting fitness consequences for offspring, but these effects are often difficult to disentangle from associated responses in offspring traits. We studied persistent maternal effects on offspring survival in North American red squirrels (Tamiasciurus hudsonicus) by manipulating maternal nutrition without altering the post-emergent nutritional environment experienced by offspring. This was accomplished by providing supplemental food to reproductive females over winter and during reproduction, but removing the supplemental food from the system prior to juvenile emergence. We then monitored juvenile dispersal, settlement and survival from birth to 1 year of age. Juveniles from supplemented mothers experienced persistent and magnifying survival advantages over juveniles from control mothers long after supplemental food was removed. These maternal effects on survival persisted, despite no observable effect on traits normally associated with high offspring quality, such as body size, dispersal distance or territory quality. However, supplemented mothers did provide their juveniles an early start by breeding an average of 18 days earlier than control mothers, which may explain the persistent survival advantages their juveniles experienced.     

Explaining the seasonal decline in litter size in European ground squirrels

In European ground squirrels Spermophilus citellus as in many ground squirrel species. late born litters are composed of fewer young than early born litters. Two alternative though not mutually exclusive hypotheses may explain this seasonal pattern of change in litter size. On the one hand. the production of few large young late in the season may be an adaptation to time limitations on the offspring. that have to complete growth and fattening prior to hibernation. Then one would expect a trade-off between offspring number and size as the breeding season progresses. At its extreme. this hypothesis would predict that total maternal effort should be equal independent of litter size. Alternatively. litter size may be determined by physiological limitations on the mother. in that highly constrained mothers breed later and produce smaller litters. Then one would expect reduced overall maternal effort in highly constrained mothers of smaller litters. In this case. a trade-off between litter size and offspring size would not be expected. We found that total maternal effort in terms of gestation length and the duration of lactation increased with increasing litter size. thus supporting the second hypothesis. Lactation was not terminated at natal emergence. It extended a relatively long period of time beyond the time of first litter emergence depending on litter size. During prolonged lactation. individual young of large litters made up body mass to young of small litters. As a consequence. juvenile weaning body mass was unaffected by litter size although offspring body mass at natal emergence was inversely related to litter size. This additional weight gain in young of large litters compensated for initial survival disadvantages and presumably affected fecundity at yearling age.     

Maternal food restriction delays weaning in the guinea pig, Cavia porcellus

In mammals, the duration of lactation varies much more than other life history parameters in relation to body mass, both within and between species. The causes of this variation are poorly understood and seem to result from varying conditions of mothers and young. We studied the effect of long-term maternal food restriction on litter mass at birth, duration of lactation and offspring development in the precocial guinea pig. Mild experimental food restriction during reproduction resulted in prolonged nursing behaviour of mothers. Evidence for a threshold mass at weaning was, however, equivocal. In the guinea pig, benefits of prolonged lactation prove hard to understand, because nutritional benefits are minor. Independently of maternal food regime, pups terminated suckling attempts several days after mothers ceased nursing behaviour. The time between the last nursing behaviour and the last suckling attempts was not longer in litters with higher need, that is, in litters of food-restricted females, than in litters of females fed ad libitum. Under food restriction, mothers maintained their own body mass, leading in pups to lower mass at birth, reduced postnatal growth and lower body mass at maturity. Guinea pig mothers appear to be selected to value their own condition more than that of a litter. We suggest that lengthening of the nursing period under poor conditions is a life history response primarily of precocial mammals.     

Early growth in male and female fallow deer fawns

In this paper we present data from a long-term study on earlygrowth and related variables in fallow deer fawns living inlarge enclosures. Pre-winter body mass was constantly higherand more strongly correlated to subadult body mass in malesthan in females. To find out the mechanism for this higher pre-wintermass in males, we analyzed the variation in pre-winter massin relation to sex, year, mother's body mass, age and parityof mother, birth date, birth mass, growth rate, suckling behavior,and other behaviors. Birth mass was higher for male fawns, andgestation length, birth date, and weaning date did not differbetween the sexes. Consequently, both pre- and postnatal growthwere faster in males than in females. No behavioral differenceswere found between the sexes that could explain the differencein postnatal growth. Pre-winter mass was positively relatedto mother's body mass. Heavy mothers gave birth earlier andto larger offspring who grew at a higher rate, independent ofoffspring sex. However, male fawns born to primiparous mothershad relatively lower growth than male fawns born to multiparousmothers. This was not the case for female fawns. Suckling timeafter the first 2 weeks was positively related to mother's bodymass and growth of offspring. However, no measurements of sucklingbehavior differed between male and female fawns. Our results,except for the effect of parity on male and female growth, indicatethat selection has not acted on mothers to promote faster earlygrowth in males.     

Mass changes of grey seal Halichoerus grypus pups on Sable Island: differential maternal investment reconsidered

Theory predicts that where variance in reproductive success differs between sexes, as in many sexually dimorphic species, mothers should invest more in male than in female offspring. In pinnipeds, parental investment is the sole responsibility of the mother. Both size at birth and subsequent mass gain during lactation are measures of that investment. We studied mass gain of grey seal pups, a sexually dimorphic species, during lactation and mass loss during the first 13 days of the postweaning fast using longitudinal data obtained on Sable Island, Canada. A mixed-effects repeated measures model was used to account for the serial correlation in these longitudinal data. We found no significant difference in the birth mass of male (16.6 ± 0.45 kg) and female (15.6 ±0.74 kg) pups. Rate of mass gain by males (3.0 ± 0.13 kg/d) and females (2.7 ± 0.12 kg/d) also did not differ significantly during the lactation period. Grey seal pups on Sable Island tended to be larger at birth and gain mass more rapidly (70%–100%) during lactation than those in the United Kingdom. Mass loss of weaned pups was curvilinear over time. Over the first 10 days of the fast, pups lost 22% of their weaning mass. In both males and females, large pups lost mass more slowly than small ones during the postweaning fast. Based on our results and a review of previous studies, we found little evidence of differential maternal investment in male and female grey seal pups.     

Sex-biased maternal investment in voles: importance of environmental conditions

Adaptive bias in sex allocation is traditionally proposed to be related to the condition of mothers as well as to the unequal fitness values of produced sexes. A positive relationship between mother condition and investment into male offspring is often predicted. This relationship was also recently found to depend on environmental conditions. We studied these causalities experimentally using a design where winter food supply was manipulated in eight outdoor-enclosed populations of field voles Microtus agrestis. At the beginning of the breeding season in spring, food-supplemented mothers seemed to be in a similar condition, measured as body mass, head width, body condition index and parasite load (blood parasite Trypanosoma), to non-supplemented mothers. Food supplements affected neither the litter size, the reproductive effort of mothers, nor the litter sex ratios at birth. However, food supplementation significantly increased the birth size of male offspring and improved their condition, as indicated by reduced parasite loads (intestinal Eimeria). Interestingly, mothers in good body condition produced larger male offspring only when environmental conditions were improved by food supplements. Although the adaptiveness of variation in mammalian sex ratios is still questionable, our study indicates that mothers in good condition bias their investment towards male offspring, but only when environmental conditions are favourable.     

根田鼠母体捕食应激效应与Trivers-Willard 模型的关系

本研究了根田鼠母体捕食应激对其子代出生、断乳和成体体重、窝性比及死亡率的作用,检验Trivers—Willard模型的2个前提条件、母体应激激素在母体投资中的作用,以及母体捕食应激效应与该模型的关系。将妊娠根田鼠母体暴露于其捕食艾鼬,母体应激子代的出生和断乳体重均显降低;到成体,雄性体重有此效应,而雌性体重则接近对照。出生窝性比无变化,但成体窝性比向雌性偏斜。不同年龄阶段的死亡率无显变化,但累计死亡率明显增大。处理雄性子代在断乳和成体时的皮质酮含量显增高,而雌性子代则无显变化,从而验证了Trivers-Willard模型的2个前提条件,提出应激母体激素参与母体对子代的投资观点,并认为,母体捕食应激使根田鼠子代成体窝性比向雌性偏斜的生理投资符合进化稳定对策。     

Competitive interactions shape offspring performance in relation to seasonal timing of emergence in Atlantic salmon

1.?Timing of birth/hatching may have strong effects on offspring fitness. Breeding time is generally considered to have evolved to match offspring arrival with optimal seasonal environmental conditions, though this is rarely tested experimentally and factors shaping the relations between timing of birth and reproductive success are often poorly understood. 2.?By manipulating incubation temperature of Atlantic salmon embryos, and hence controlling for maternal and genetic effects, we obtained offspring emerging from nests prior to (accelerated), during and after (decelerated) normal emergence times, and accordingly experienced widely different seasonal environmental conditions at emergence (stream temperature range 4-16 °C). The accelerated group emerged at temperatures that are generally considered to be highly sub-optimal for growth and likely imposes strong constraints on feeding and activity, and during a peak in water discharge which is expected to negatively influence habitat availability. 3.?In the wild, overall mortality during the period after emergence was 79%, and was significantly affected by both release density and emergence timing. Accelerated offspring, which emerged earliest and experienced the harshest environmental conditions, had both highest survival and largest final body size. The effect was particularly strong at the high density release site, where survival of accelerated offspring was significantly higher than both the normal and decelerated groups. 4.?In more controlled semi-natural environments, all developmental groups were able to perform well, but accelerated offspring had a relatively better performance than the later emerging offspring when density was high. Furthermore, the relative performance of the different groups was not sensitive to water discharge regimes (temporally stable vs. fluctuating). 5.?These results suggest that the performance of offspring in relation to seasonal timing of emergence is highly affected by competitive interactions in Atlantic salmon. Although a match between phenology and optimal seasonal environmental conditions may be highly important for organisms depending on specific resources that are only available during a limited period of the season, such resources may be available in variable amounts year around for many organisms. For these, offspring success may to a larger degree be shaped by the timing of their hatching/birth relative to each other, and particularly so under high population densities.     

Growth and survivorship in juvenile woodlice: is birth mass important?

Growth and survivorship of individually reared offspring of the woodlice (Isopoda Oniscidae) Porcellionides pruinosus (Brandt) and Aphiloscia vilis (Budde-Lund) were measured under laboratory conditions These measures of offspring fitness were compared to birth mass, which is a realistic measure of parental investment in offspring for these species, to test a key assumption of life history theory that an increase in resource allocation to offspring will increase offspring fitness In both species birth mass had no significant effect on survivorship to 60 days and in P pruinosus birth mass did not correlate with mass at 15, 30 or 45 days after birth In A vilis larger young at birth had a slight growth advantage to 15 days but not thereafter There appears to be little effect of birth mass on offspring fitness despite a considerable range in birth mass within broods In these species high variance in juvenile growth may spread subsequent reproductive effort and avoid risks of total reproductive failure     

Causes and consequences of variation in offspring body mass: meta‐analyses in birds and mammals

Early survival is highly variable and strongly influences observed population growth rates in most vertebrate populations. One of the major potential drivers of survival variation among juveniles is body mass. Heavy juveniles are better fed and have greater body reserves, and are thus assumed to survive better than light individuals. In spite of this, some studies have failed to detect an influence of body mass on offspring survival, questioning whether offspring body mass does indeed consistently influence juvenile survival, or whether this occurs in particular species/environments. Furthermore, the causes for variation in offspring mass are poorly understood, although maternal mass has often been reported to play a crucial role. To understand why offspring differ in body mass, and how this influences juvenile survival, we performed phylogenetically corrected meta‐analyses of both the relationship between offspring body mass and offspring survival in birds and mammals and the relationship between maternal mass and offspring mass in mammals. We found strong support for an overall positive effect of offspring body mass on survival, with a more pronounced influence in mammals than in birds. An increase of one standard deviation of body mass increased the odds of offspring survival by 71% in mammals and by 44% in birds. A cost of being too fat in birds in terms of flight performance might explain why body mass is a less reliable predictor of offspring survival in birds. We then looked for moderators explaining the among‐study differences reported in the intensity of this relationship. Surprisingly, sex did not influence the intensity of the offspring mass–survival relationship and phylogeny only accounted for a small proportion of observed variation in the intensity of that relationship. Among the potential factors that might affect the relationship between mass and survival in juveniles, only environmental conditions was influential in mammals. Offspring survival was most strongly influenced by body mass in captive populations and wild populations in the absence of predation. We also found support for the expected positive effect of maternal mass on offspring mass in mammals (r_pearson = 0.387). As body mass is a strong predictor of early survival, we expected heavier mothers to allocate more to their offspring, leading them to be heavier and so to have a higher survival. However, none of the potential factors we tested for variation in the maternal mass–offspring mass relationship had a detectable influence. Further studies should focus on linking these two relationships to determine whether a strong effect of offspring size on early survival is associated with a high correlation coefficient between maternal mass and offspring mass.     

Factor affecting body mass of young fat dormics (Glis glis) at weaning and by hibernation

This paper describes the factors affecting body mass at weaning and by hibernation of young fat dormice (Glis glis) We studied a free-living population in 1991 and 1992 in a mixed forest where 100 nest-boxes were placed. In the two years, 128 females used the nest-boxes to give birth and rear their litters and more than 900 dormice have been individually marked. Body mass of young at weaning was positively correlated to body mass of the mother in both years and to birth date in 1991, while litter size was negatively correlated to body mass of the young in 1992 only, when birth took place about two weeks later than in 1991. On the contrary, by the time of hibernation, young born later weighed significantly less than those born earlier in the season, and heavier young at weaning remained heavier by hibernation, while maternal body mass and litter size did not significantly affect body mass of young at this stage. Body mass of the mother was negatively related to the date of parturition and early breeding females lost weight during lactation, while females that bred later did not. It seems, therefore, that early breeding carries a cost that only females in good condition can afford. It appears that females may follow two strategies: those in poor body condition will delay pregnancy so that lactation occurs when environmental feeding conditions are at their best; those in good condition will breed earlier, since they can afford to lose weight during lactation. Their young will have more time to grow before hibernation and the mothers themselves will have a longer period of time for restoring their fat reserves.     

Sex-specific fitness returns are too weak to select for non-random patterns of sex allocation in a viviparous snake

When environmental conditions exert sex-specific selection on offspring, mothers should benefit from biasing their sex allocation towards the sex with the highest fitness in a given environment. Yet, studies show mixed support for such adaptive strategies in vertebrates, which may be due to mechanistic constraints and/or weak selection on facultative sex allocation. In an attempt to disentangle these alternatives, we quantified sex-specific fitness returns and sex allocation (sex ratio and sex-specific mass at birth) according to maternal factors (body size, age, birth date, and litter size), habitat, and year in a viviparous snake with genotypic sex determination. We used data on 106 litters from 19 years of field survey in two nearby habitats occupied by the meadow viper Vipera ursinii ursinii in south-eastern France. Maternal reproductive investment and habitat quality had no differential effects on the growth and survival of sons and daughters. Sex ratio at birth was balanced despite a slight female-biased mortality before birth. No sexual mass dimorphism between offspring was evident. Sex allocation was almost random apart for a trend towards more male-biased litters as females grew older, which could be explained by an inbreeding avoidance strategy. Thus, a weak selection for facultative sex allocation seems sufficient to explain the almost equal sex allocation in the meadow viper.     

Reception of photoperiodic information by fetal Siberian hamsters: role of the mother's pineal gland

The rate of reproductive development in juvenile male Siberian hamsters is strongly influenced by daylength (photoperiod). Recent studies indicate that reception of photoperiodic cues begins during fetal life. The present experiments yielded a further demonstration that developing male Siberian hamsters receive information about the photoperiod to which their mother is exposed during pregnancy. The possibility that photoperiodic information is transmitted from mother to young after birth was investigated by cross-fostering young gestated on 12L and 16L to mothers from the other photoperiod. Litters were cross-fostered on the day of birth and then were transferred, along with their foster mothers, to 14L. We found no influence of the mother after birth, indicating that transmission of photoperiodic information from mother to young must occur during gestation. To determine if the pineal gland of the mother is required for this response, adult females were pinealectomized or sham-operated and paired with intact males in 12L, 14L, or 16L. After parturition parents and offspring were exposed to 14L. The influence of prenatal photoperiod on postnatal testicular development in 14L was blocked by pinealectomy of the mother. Postnatal testicular development was retarded in offspring that experienced a photoperiod transfer from either 15L to 14L or 8L to 12L at birth. In contrast, the inhibitory effect of a transfer from 16L to 14L at birth was abolished when juvenile males were exposed to a single long photoperiod (16.3 h light) at age 17-21 days and then were returned to 14L.     

Juvenile exposure to predator cues induces a larger egg size in fish

When females anticipate a hazardous environment for their offspring, they can increase offspring survival by producing larger young. Early environmental experience determines egg size in different animal taxa. We predicted that a higher perceived predation risk by juveniles would cause an increase in the sizes of eggs that they produce as adults. To test this, we exposed juveniles of the mouthbrooding cichlid Eretmodus cyanostictus in a split-brood experiment either to cues of a natural predator or to a control situation. After maturation, females that had been confronted with predators produced heavier eggs, whereas clutch size itself was not affected by the treatment. This effect cannot be explained by a differential female body size because the predator treatment did not influence growth trajectories. The observed increase of egg mass is likely to be adaptive, as heavier eggs gave rise to larger young and in fish, juvenile predation risk drops sharply with increasing body size. This study provides the first evidence that predator cues perceived by females early in life positively affect egg mass, suggesting that these cues allow her to predict the predation risk for her offspring.     

Growth and mortality in juvenile European wild rabbits (Oryctolagus cuniclus)

Factors influencing the growth and mortality patterns of young were examined in a small population of European wild rabbits. Juveniles were trapped and tagged for individual identification soon after their emergence from underground nests. Age was estimated from weight on first capture and survivorship determined from the last date individuals were observed. Differences in growth rates of young born throughout the season and between the three years of the study were related to population density. Maternal social status had no significant effect on post-emergence growth rates of kittens, or on offspring survival. Young born in females' first litters of the season, in March, had significantly lower mortality than subsequent litters. Cats and stoats were regularly observed on the study site and are likely to have been major agents of juvenile mortality. There was no difference in the growth rates of the sexes, and correspondingly non-breeding adult males and females were of similar body weight. From their first days above ground juvenile males suffered significantly higher mortality than females. Differential mortality may be linked to previously reported sex differences in behaviour if the greater activity of young males made them more vulnerable to predation or attack by neighbouring territorial conspecifics.     

Reactions of nuclear-family–reared rhesus macaques to the births of younger siblings

A group of juvenile rhesus macaques (Macaca mulatta) living in a nuclear-family laboratory environment was studied to determine their responses to the births of siblings. The frequencies of interactions with family members (mothers, fathers, and new siblings) and nonfamily (peers, unrelated infants, and unrelated adults) were studied over both the year preceding and the year following sibling birth. The frequencies of specific behaviors in each of those interactions and the frequencies of interactions in each area of the nuclear-family unit (home cage, play area, or other families' cage) were also examined. After new siblings arrived, several measures of interactions with mothers, fathers, and new siblings increased significantly; by contrast interactions with peers decreased substantially over the post-birth year. Although the frequency of interactions in home cages remained stable over the 2-year period, interactions outside of the subjects' home cages decreased significantly after siblings were born. An additional subject group whose mothers became pregnant but failed to deliver viable offspring showed no significant changes in total levels of interactions with peers; they did, however, exhibit increases in some interactions with unrelated infants and adults. Female juveniles interacted with new siblings significantly more often than did males when siblings were less than 6 months old, but as siblings grew older (6–12 months), females' levels of interaction with them fell to a level equal to that of males. In the nuclear-family social structure, the birth of a sibling resulted in an increased emphasis on family interactions at the expense of peer interactions.     

Survival and growth of developing rats during centrifugation at 2G

We studied the effects of 2G hypergravity on the survival, body mass and growth of postnatal rats (Rattus norvegicus). Nursing litters comprised of either neonatal (Postnatal day [P]7) or pre-weanling (P14) rats and their mothers were exposed to 16 days of continuous centrifugation. All of the offspring survived and gained body mass, indicating that mothers nursed their young. Following the onset of centrifugation, neonatal and pre-weanling rats showed a reduction in growth relative to age-matched environmental controls (EC). At the completion of testing, body mass of the hypergravity (HG) groups was significantly less than that of controls (p<0.05). Over the course of the test, the HG-exposed P7 group showed an overall 55% gain in body mass as compared to a 71% increase in controls, while the HG-exposed P14 group showed a 62% increase relative to 75% in controls. Neonatal offspring (P7) gained body mass during centrifugation, but at significantly slower rates as compared to EC controls (p<0.05). In contrast, growth rates of pre-weanling (P14) rats were not reduced relative to controls, possibly related to the initiation of weaning, around P18 in the rat. These findings raise key issues relevant to studies of nursing mammals reared in altered gravity.     

Offspring size at weaning affects survival to recruitment and reproductive performance of primiparous gray seals

Offspring size affects survival and subsequent reproduction in many organisms. However, studies of offspring size in large mammals are often limited to effects on juveniles because of the difficulty of following individuals to maturity. We used data from a long‐term study of individually marked gray seals (Halichoerus grypus; Fabricius, 1791) to test the hypothesis that larger offspring have higher survival to recruitment and are larger and more successful primiparous mothers than smaller offspring. Between 1998 and 2002, 1182 newly weaned female pups were branded with unique permanent marks on Sable Island, Canada. Each year through 2012, all branded females returning to the breeding colony were identified in weekly censuses and a subset were captured and measured. Females that survived were significantly longer offspring than those not sighted, indicating size‐selective mortality between weaning and recruitment. The probability of female survival to recruitment varied among cohorts and increased nonlinearly with body mass at weaning. Beyond 51.5 kg (mean population weaning mass) weaning mass did not influence the probability of survival. The probability of female survival to recruitment increased monotonically with body length at weaning. Body length at primiparity was positively related to her body length and mass at weaning. Three‐day postpartum mass (proxy for birth mass) of firstborn pups was also positively related to body length of females when they were weaned. However, females that were longer or heavier when they were weaned did not wean heavier firstborn offspring.