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.     

Reproductive Decisions in Female European Ground Squirrels: Factors Affecting Reproductive Output and Maternal Investment

Physiological and behavioural parameters associated with reproductive effort and success were investigated in female European ground squirrels Spermophilus citellus . The proportion of reproductive (lactating) females in the study population was over 90% and was not related to age. Timing of oestrus and ovulation was found to be affected by the female's emergence date and condition. Females with low emergence mass showed delayed oestrus. Differences in ovulation dates were shown to affect reproductive output in terms of litter size and sex ratio. Early litters were larger and male biased. X-ray techniques were used to determine intrauterine litter size in individual females. The results indicated that litter size and sex ratio were fixed prenatally. Lactation costs were reflected in the intensity of mass loss and duration of lactation. Mass loss varied with litter size, in that females with large litters showed a more rapid loss than others. The second parental investment parameter, lactation duration, varied among individual females and was dependent on the timing of reproduction and litter size (except yearlings). Early born litters, which were, in most cases, larger than later ones, were nursed longer. Prolonged lactation periods affected female condition in that they started prehibernation fattening later and entered hibernation with a lower mass than individuals that had shorter lactation periods. Yearling females probably could not afford the energetic costs of long lactation, independent of their offspring number. These results indicated that females with higher reproductive output and higher investment were unable to compensate these costs before hibernation. Consequences for these individuals could therefore be lower over-winter survival or a delayed oestrus in the following season.     

Reproductive effort during gestation and lactation by Richardson's ground squirrels

Summary Reproductive effort by yearling and older female Richardson's ground squirrels was studied over a 4-year period in southern Alberta by obtaining serial weight records from marked individuals to compare the mother's mass at critical points in the annual cycle (emergence from hibernation, estrus, parturition, and litter emergence) with her litter's mass at birth and weaning. Yearlings weighed only 80% of older adults at emergence from hibernation, but they mated at the same time as older females, attained adult mass coincident with pregnancy, and weaned litters that were not significantly smaller in size or mass than those of older females. Age and maternal mass were weak predictors of litter size and litter mass. Of the net increase in mass of the combined mother-litter unit during gestation, over half (60% of 139 g for yearlings; 52% of 127 g for older females) was attributable to an increase in the mother's own mass, whereas during lactation almost all of the net increase (93% of 545 g for yearlings; 96% of 567 g for older females) was attributable to an increase in the litter's mass. On a daily basis, deposition of mass in the litter was 6 times greater during lactation than gestation. On average, neonates weighed 2.3% (6.5 g) of maternal mass at birth and 23.1% (81 g) at emergence from the natal burrow; offspring masses at birth and at emergence were significantly negatively correlated with litter size. On average, litters weighed 16.3% (48 g) of maternal mass at birth and 157.5% (578 g) at emergence from the natal burrow. Compared with other hibernating sciurids, Richardson's ground squirrels have a similar offspring mass relative to maternal mass both at birth and at emergence from the natal burrow. However, because of the large litter size (typically 6–8), absolute reproductive effort, measured either as litter mass at birth or at natal emergence, is large for the body size of the species.     

Body size,litter size,timing of reproduction,and juvenile survival in the Unita ground squirrel,Spermophilus armatus

The timing of reproduction affected litter size, offspring mass, and offspring survival in the Uinta ground squirrel, Spermophilus armatus, in Grand Teton National Park, Wyoming. Survival of juvenile females to yearling age varied negatively with date of weaning and positively with individual offspring mass. At the same time, juveniles weaned early in the season were lighter, and juveniles weaned later in the season were heavier. The coefficient of variation for juvenile body mass, originally measured at weaning, significantly decreased by the time juveniles entered hibernation, indicating that individuals weaned early and light caught up in body mass to individuals weaned later and heavier. From the perspective of the mother's investment in the litter, litter size (corrected for mother's mass) decreased with later wcaning dates, while the relationship of weaning date to litter mass (corrected for mother's mass) was significant in only one year. Maternal allocation of resources in litters changed over the season so that mothers produced many, small offspring early in the season, and fewer, large offspring late in the season.     

Parasite removal and its impact on litter size and body condition in Columbian ground squirrels (Spermophilus columbianus)

The current interest in studies of parasite-host relationships is focusing on the impact of parasites on mate choice, sexual selection and individual fitness. While most studies have been done on birds and fishes, little is known about the interactions between parasites and reproductive success in mammals. In this study, I experimentally removed ectoparasites (mainly fleas) from some female Columbian ground squirrels using a commercially available powder, whereas other squirrels were left untreated as controls. The anti-parasite treatment led to an increase in female body condition during lactation and at weaning and an increase in litter size. The increase of body mass in treated females between birth of the young and weaning is also different from observations done over 8 years in the same population, where untreated females usually lost mass. In close to 400 litters from untreated females in the same area, only 2% had litter sizes as big as the treated females. I conclude that ectoparasites can have a profound impact not only on individuals but probably on the dynamics of the whole population.     

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.     

Influences of the feeding ecology on body mass and possible implications for reproduction in the edible dormouse (<Emphasis Type="Italic">Glis glis</Emphasis>)

The edible dormouse (Glis glis) is a small rodent and an obligate hibernator. Dormice undergo strong fluctuations of reproductive output during years that seem to be timed to coincide with future food supply. This behaviour enables them to avoid producing young that will starve with a high probability due to food shortage, and to increase their lifetime reproductive success. Aims of this study were to elucidate the extent to which feeding ecology in the edible dormouse has an impact on body mass and the fatty acid (FA) pattern of the white adipose tissue (WAT) before and after hibernation, which in turn might influence reproductive status in spring. Dormice show strong seasonal fluctuations of the body mass, which is reduced by one third during hibernation. Body mass and its changes depend on autumnal food availability as well as on the dietary FA pattern. During the pre-hibernation fattening period, dormice eat lipid rich food with a high content of linoleic acid. During hibernation, linoleic acid content is slightly but significantly reduced and body mass loss during winter is negatively correlated with the pre-hibernation linoleic acid content in the WAT. No relation between reproductive status and body mass, body condition or the FAs pattern of the WAT could be detected. However, in a year of high reproduction, dormice commence the shift to seed eating earlier than in a year of low reproduction. These seeds could be either a predictor for future food supply in autumn, or represent a high-energy food compensating high energetic costs of sexual activity in male edible dormice.     

Polyphasic growth-weight response in the guinea pig

An assessment of the growth rate pattern based on longitudinal weight data of guinea pigs was conducted over the presumptive preweaning and early postweaning period. The study covered the minimum of 44 and maximum of 53 days with means calculated for 3-day intervals involving 13 litters and a total of 40 young. Initial analysis showed males heavier than females at birth and both sexes being inversely proportional to litter size at birth. At the end of the experimental period the males were still heavier than the females but their weight distribution peaked in litter of three while the females were heaviest in the largest litters of five and lightest in the litter of one. Phasic linear regression line analysis revealed 4 litters with a triphasic growth outline and 6 litters with quadrophasic growth outline. Final assessments indicate that the total period of 53 days manifests a quadrophasic growth outline while, in contrast, the period specifically involved with the process of lactation and weaning displays a definite diphasic growth spectrum and is comparable to the tri- or quadrophasic spectrum occurring in the albino rats, rice rats and hamsters.     

黑线仓鼠繁殖输出与基础代谢率的关系

为了解黑线仓鼠繁殖输出与基础代谢率(BMR)的关系,阐明最大持续能量收支(SusMR)的限制水平, 揭示哺乳期能量收支对策,本文测定了哺乳期黑线仓鼠的体重、摄食量、BMR 和身体组成,以及哺乳期的胎仔数、胎仔重和泌乳能量支出(MEO)。结果显示,黑线仓鼠哺乳期体重降低了15.0 ± 0.8% , 摄食量显著增加, 哺乳高峰期平均摄食量为13.9 ± 0.3 g /d, 摄入能为222.1 ± 5.3 kJ/ d, 比哺乳初期增加121% , 比对照组高288% ;哺乳高峰期MEO 为62.4 ± 2.3 kJ/ d, 哺乳末期BMR 为49.7 ± 1.1 kJ/ d; 断乳时平均胎仔数4.7 ± 0.2、窝胎仔重50.5 ±1.6 g; 哺乳末期BMR 比对照组增加48% ,BMR 与消化系统各器官的相关性高于对照组; BMR 与胎仔数、胎仔重、乳腺重量和MEO 显著正相关。结果表明:初次繁殖的黑线仓鼠哺乳期SusMR 限制为4.47 ×BMR, 在自身维持和繁殖输出之间采取了“权衡分配”的原则,通过体重降低以减少BMR 的增加幅度, 从而有利于繁殖输出。     

Reproductive performance in grey seals: age-related improvement and senescence in a capital breeder

1. Three hypotheses have been advanced to account for age-related improvement in performance: the selection hypothesis predicts improved due to the loss of lower quality phenotypes, the constraint hypothesis predicts individuals improve function, and the restraint hypothesis predicts younger individuals forego or reduce effort because of mortality risks. A decline in age-related performance (i.e. senescence) is predicted by mutation accumulation, antagonistic pleiotropy and disposable soma (wear and tear) hypotheses. 2. Using five measures of performance - birth rate, maternal and pup birth mass, pup weaning mass, weaning success and lactation length - we tested these hypotheses concerning age-related change in reproduction in 279 female grey seals (Halichoerus grypus), ages 4-42 years, over a 23-year period between 1983 and 2005 on Sable Island, Nova Scotia. These females produced 2071 pups. 3. Although body mass of primiparous females increased with age (4-7 years) birth mass of their pups did not, but pup weaning mass did. Second- and third-parity females of the same age as primiparous females gave birth to and weaned heavier pups. However, parity and age were dropped from models when maternal body mass was included. 4. The proportion of females giving birth varied significantly with maternal age, increasing in young females and then declining late in life. Weaning success rate also increased rapidly to about 8 years and subsequently declined in females > 32 years. 5. Generalized additive models indicated nonlinear changes in 3 day body mass (i.e. approximately birth mass) and weaning mass of pups as a function of maternal age, after accounting statistically for the effects of maternal body mass. Mixed-effects, repeated-measures models fitted to longitudinal data further supported the conclusion that pup birth mass and weaning mass vary nonlinearly with maternal age and indicated nonlinear changes in lactation duration. 6. We found some support for the constraint hypothesis, but our findings were not consistent with the selection hypothesis or the restraint hypothesis as the basis for improvement in reproductive performance. 7. Senescence was evident in multiple female and offspring traits, indicating the degeneration in function of several physiological systems as predicted by the disposable soma hypothesis.     

Factors Affecting Date of Implantation,Parturition, and Den Entry Estimated from Activity and Body Temperature in Free-Ranging Brown Bears

Knowledge of factors influencing the timing of reproduction is important for animal conservation and management. Brown bears (Ursus arctos) are able to vary the birth date of their cubs in response to their fat stores, but little information is available about the timing of implantation and parturition in free-ranging brown bears. Body temperature and activity of pregnant brown bears is higher during the gestation period than during the rest of hibernation and drops at parturition. We compared mean daily body temperature and activity levels of pregnant and nonpregnant females during preimplantation, gestation, and lactation. Additionally we tested whether age, litter size, primiparity, environmental conditions, and the start of hibernation influence the timing of parturition. The mean date of implantation was 1 December (SD = 12), the mean date of parturition was 26 January (SD = 12), and the mean duration of the gestation period was 56 days (SD = 2). The body temperature of pregnant females was higher during the gestation and lactation periods than that of nonpregnant bears. The body temperature of pregnant females decreased during the gestation period. Activity recordings were also used to determine the date of parturition. The parturition dates calculated with activity and body temperature data did not differ significantly and were the same in 50% of the females. Older females started hibernation earlier. The start of hibernation was earlier during years with favorable environmental conditions. Dates of parturition were later during years with good environmental conditions which was unexpected. We suggest that free-ranging pregnant brown bears in areas with high levels of human activities at the beginning of the denning period, as in our study area, might prioritize investing energy in early denning than in early parturition during years with favorable environmental conditions, as a strategy to prevent disturbances caused by human.     

Variations in adult body mass in roe deer: the effects of population density at birth and of habitat quality

Body mass is a key determinant of fitness components in many organisms, and adult mass varies considerably among individuals within populations. These variations have several causes, involve temporal and spatial factors, and are not yet well understood. We use long-term data from 20 roe deer cohorts (1977-96) in a 2600 ha study area (Chizé, western France) with two habitats contrasting in quality (rich oak forest in the North versus poor beech forest in the South) to analyse the effects of both cohort and habitat quality on adult mass (i.e. median body mass between 4 and 10 years of age) of roe deer (Capreolus capreolus). Cohort strongly influenced the adult body mass of roe deer in both sexes: males born in 1994 were 5.2 kg heavier when aged between 4 and 10 years old than males born in 1986, while females born in 1995 were 4.7 kg heavier between 4 and 10 years old than females born in 1982. For a given cohort, adult males were, on average, 0.9 kg heavier in the rich oak forest than in the poor beech forest. A similar trend occurred for adult females (0.5 kg heavier in the oak forest). The effects of cohort and habitat were additive and accounted for ca. 40% of the variation observed in the adult mass of roe deer at Chizé (males: 41.2%; females: 40.2%). Population density during the spring of the birth accounted for about 35% of cohort variation, whereas rainfall in May-June had no effect. Such delayed effects of density at birth on adult body mass probably affect population dynamics, and might constitute a mechanism by which delayed density-dependence occurs in ungulate populations.     

Body development in sows, feed intake and maternal capacity. Part 2: gilt body condition before and after lactation, reproductive performance and correlations with lactation feed intake

Data on sow body weight (BW) and fatness (n = ~2250 pregnant sows) and reproductive data (including historical: n = ~18 000) were used to examine the genetic and phenotypic associations between body condition before and after farrowing, gestational outcomes, lactation feed intake and the gilts' ability to survive unculled to farrow in the second parity. Within-trait genetic correlations were very high between weight (0.77 ± 0.06) and fat depth (0.91 ± 0.04) recorded before farrowing and at weaning. Litter size traits were generally uncorrelated genetically with aspects of sow BW and body condition. However, genetic correlations indicated that sows producing heavier piglets at birth had litters with increased gain (0.36 ± 0.16), and were characterised by greater weight (-0.72 ± 0.08) and fat change (-0.19 ± 0.15) during lactation, reflected to a lesser extent by lower weight (-0.12 ± 0.11) and fatness (-0.17 ± 0.10) at weaning. Genetic correlations (r(a)) between reproductive traits and lactation feed intake were generally low, but favourable. However, lactation intake was positively correlated with measures of sow size (r(a) = ~0.55), such that selection for lactation feed intake would likely be accompanied by increased mature sow size. Phenotypic correlations (r(p)) showed that sow survival to the second parity (FAR12) was positively influenced by litter size and fat depth at weaning, supporting attributes of increased fatness before farrowing, less weight loss during lactation and an increased lactation intake.     

Litter sex ratios in Richardson’s ground squirrels: long-term data support random sex allocation and homeostasis

When costs of producing male versus female offspring differ, parents may vary allocation of resources between sons and daughters. We tested leading sex-allocation theories using an information-theoretic approach and Bayesian hierarchical models to analyse litter sex ratios (proportion males) at weaning for 1,049 litters over 24 years from a population of Richardson’s ground squirrels (Urocitellus richardsonii), a polygynandrous, annually reproducing mammal in which litter size averages from six to seven offspring and sons are significantly heavier than daughters at birth and weaning. The model representing random Mendelian sex-chromosome assortment fit the data best; a homeostatic model received similar support but other models performed poorly. Embryo resorption was rare, and 5 years of litter data in a second population revealed no differences in litter size or litter sex ratio between birth and weaning, suggesting that litter size and sex ratio are determined in early pregnancy. Sex ratio did not vary with litter size at weaning in any of 29 years, and the observed distribution of sex ratios did not differ significantly from the binomial distribution for any litter size. For 1,580 weaned litters in the two populations, average sex ratio deviated from parity in only 3 of 29 years. Heavier females made a greater reproductive investment than lighter females, weaning larger and heavier litters composed of smaller sons and daughters, but litter sex ratio was positively related to maternal mass in only 2 of 29 years. Such occasional significant patterns emphasize the importance of multi-season studies in distinguishing infrequent events from normal patterns.     

Age-related reproductive success in solitarily and communally nesting female dormice (Glis glis)

The factors underlying the occurrence of communal nesting in a natural population of fat dormouse ( Glis glis ) breeding in nest-boxes have been analysed on the assumption that such behaviour is adaptive. Since co-nesting females were previously shown to be close kin (prevalently mother-daughter pairs), they were expected to obtain kin-selected benefits not enjoyed by solitary breeders. The only advantage so far detected accrues to the younger partner, which gives birth, on average, several days earlier than it would as a solitary breeder and thus allows its progeny to attain a heavier weight by the time of hibernation. On the other hand, pup predation in the nest, presumably caused by the older female towards the younger's litter, was occasionally recorded. On the whole, it is perhaps more plausible that communal nesting results from delayed dispersal of yearling females when a high population density limits breeding space, as proposed for other small rodents.     

Annual cycle of energy and time expenditure in a golden-mantled ground squirrel population

Summary We have analyzed seasonal shifts of energy and time allocation in a population of golden-mantled ground squirrels (Spermophilus saturatus) by directly measuring total daily energy expenditure (DEE) with an isotopic technique (doubly labeled water=dlw), and by estimating components of total DEE through an integration of field behavioral observations with laboratory-measured rates of energy expenditure (oxygen consumption) associated with major behavioral and physiological states. Hibernation laster about 7 1/2 months, and the 4 1/2-month activity season consisted of mating, a 28-d gestation of 3–5 young, 5 1/2 weeks of postnatal growth building to a peak in lactation just before the young emerged above ground, an additional 2–3-week period of maternal care before dispersal, and finally restoration of body mass preceding hibernation. Although the hibernation season comprised nearly two-thirds of the year, it involved only 13–17% of annual energy expenditure, leaving about 85% of energy expenditure for the active season. Ground squirrels were actually present on the surface for only about 11% of the year's time, and the foraging time required to obtain the total annual energy supply amounted to only about 2% of the year's time. The squirrels fed mainly on herbs in the early season and hypogeous fungi later; both were used extensively during peak lactation when female energy expenditure and demand were maximal. Average daily foraging time increased steadily throughout the season to a maximum of 28% of aboveground time as availability of greens diminished and fungus predominated in the diet; time availability did not limit foraging since the animals sat on average for 65% of the daily surface time of about 7 h. Timing of reproduction is apparently optimized such that peak reproductive energy demands are matched with maximal food availability and moderate thermal conditions that minimize energy demand. Despite the greater body mass of males, the greatest total DEE (measured by dlw) of any squirrels at any time of year was that of females during peak lactation. For production of young and lactation through above-ground emergence of an average litter of 2.7, females required a total energy increase of 24% above annual nonreproductive metabolism. Yearling females all bred and performed similarly to older females, yet some costs were greater because the yearlings began and ended hibernation at smaller mass, compensated by giving birth later, and finally showed a greater absolute increase in body mass over the active season than older females. Annual metabolic energy expenditure of breeding males was about 18% greater than that of females, due to greater male body mass. Yet the annual energy intake requirement for both sexes was essentially identical (about 42MJ) due to the greater reproductive export by females in the form of newborn and milk. During the mating season males showed wide-ranging exploratory behavior and social interactions, including aggression, that involved considerable locomotory energy expenditures. Although we did not directly account for the energetics of these specific reproductive behaviors, they are critical to male reproductive success and on a daily basis they probably involved much greater energy expenditure than sperm production. Some yearling males avoided these costs by foregoing testicular development, yet they allocated four times as much energy to growth as older males, thereby increasing somatic condition for the future.     

Kin Recognition and Maternal Care under Restricted Feeding in House Mice (Mus domesticus)

This study tests the hypothesis that female house mice (F1 generation of wild caught Mus domesticus) should preferentially invest in own offspring if confronted with young of different degrees of relatedness. The maternal behaviour of females with litters of 4 own and 4 unrelated alien young (cross-fostered at day 1 of lactation) was analysed during a lactation period of 22 days both under ad libitum and under restricted feeding (food was restricted by 20%). Cross-fostering and restricted feeding had no effect on the amount of time spent nursing until weaning. Under both feeding conditions the females did not differ in their maternal behaviour towards own and alien young: there were no significant differences either in the amount of time spent nursing own versus alien pups or in the time spent licking own versus alien young. Weight gain of own and alien = wild littermates did not differ significantly in mixed litters and was similar both under ad libitum and under restricted feeding. Such indiscriminate behaviour might be adaptive if female house mice prefer to communally nest with a relative and thus improve their inclusive fitness by investing in own and related offspring in a communal nest. Under moderate restricted feeding females could not wean the entire litter but reduced litter size by cannibalizing on average 2.7 pups (75% of the pups were killed when they were 4–8 days old). Females with cross-fostered litters killed as many own as alien young. This suggests that females cannot discriminate between own and unrelated young if cross-fostering takes place at day 1 of lactation. Besides testing kin recognition abilities, the experiments also allow analysis of the weaning strategy of females under food shortage. Under restricted feeding, body weight of the females was significantly lower during middle lactation than under ad libitum feeding. Weaning weight of young in reduced litters under food restriction (9–10 g) did not differ significantly from weaning weight of young in litters of 7–10 young, but was lower than that of young in similar sized litters (litter size 6), under ad libitum feeding. The maternal behaviour of cannibalizing some young under food shortage can be interpreted as a weaning strategy which results in the largest number of offspring that can be raised to a minimal weaning weight of 9–10 g. Such a weaning strategy might represent a favourable trade-off between number and size of young produced.     

Influence of maternal mass and condition on energy transfer in Weddell seals

1. Environmental variation influences food abundance and availability, which is reflected in the reproductive success of top predators. We examined maternal expenditure, offspring mass and condition for Weddell seals in 2 years when individuals exhibited marked differences in these traits. 2. For females weighing > or = 355 kg there was a positive relationship between maternal post-partum mass (MPPM) and lactation length, but below this there was no relationship, suggesting that heavier females were able to increase lactation length but lighter females were restricted to a minimum lactation period of 33 days. 3. Overall, females were heavier in 2002, but in 2003 shorter females were lighter than similar-sized females in 2002 suggesting that the effects of environmental variability on foraging success and condition are more pronounced in smaller individuals. 4. There was no relationship between MPPM and pup birth mass, indicating pre-partum investment did not differ between years. However, there was a positive relationship between MPPM and pup mass gain. Mass and energy transfer efficiency were 10.2 and 5.4% higher in 2002 than 2003, which suggests costs associated with a putatively poor-resource year were delayed until lactation. 5. Heavier females lost a higher proportion of mass during lactation in both years, so smaller females may not have been able to provide more to their offspring to wean a pup of similar size to larger females. 6. MPPM had only a small influence on total body lipid; therefore, regardless of mass, females had the same relative body composition. Females with male pups lost a higher percentage of lipid than those with female pups, but by the end of lactation female pups had 4.5% higher lipid content than males. 7. It appears that for Weddell seals the consequences of environmentally induced variation in food availability are manifested in differences in maternal mass and expenditure during lactation. These differences translate to changes in pup mass and condition at weaning with potential consequences for future survival and recruitment.     

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.     

Birthdate, mass and survival in mountain goat kids: effects of maternal characteristics and forage quality

In temperate environments, early-born ungulates may enjoy a longer growth period before winter, and so attain a higher body mass and an increased probability of survival compared to late-born ones. We assessed the effects of maternal characteristics, forage quality and population density on kid birthdate, mass and survival in a population of marked mountain goats (Oreamnos americanus) in Alberta. The duration and timing of the birth season were similar in all years. Births were highly synchronised: 80% of kids were born within 2 weeks of the first birth. Maternal age, maternal social rank and density did not affect kid birthdate or mass. Previous breeding experience was not related to kid birthdate, but kids born to pluriparous mothers were heavier during summer than kids born to primiparous mothers. Male and female kids had similar mass and accumulated mass linearly during summer. Early-born kids were heavier than late-born kids. Faecal crude protein (FCP) in late spring and maternal mass were positively related to kid mass. Survival to weaning appeared higher for males (90%) than for females (78%), but survival to 1 year was 65% for both sexes. FCP in late spring, density, birthdate and mass did not affect kid survival to weaning in either sex. Survival to 1 year increased with FCP in late spring for females, but not for males. Survival to 1 year was independent of birthdate for both sexes, but heavy females survived better than light ones. Multiple logistic regression revealed a positive effect of mass on survival to 1 year when the sexes were pooled. Our results suggest that mountain goats are constrained to give birth in a short birth season synchronised with forage productivity.