Ectoparasites and fitness of female Columbian ground squirrels

Parasites play an important role in the evolution of host traits via natural selection, coevolution and sexually selected ornaments used in mate choice. These evolutionary scenarios assume fitness costs for hosts. To test this assumption, we conducted an ectoparasite removal experiment in free-living Columbian ground squirrels (Urocittelus columbianus) in four populations over three years. Adult females were randomly chosen to be either experimentally treated with anti-parasite treatments (spot-on solution and flea powder, N = 61) or a sham treatment (control, N = 44). We expected that experimental females would show better body condition, increased reproductive success and enhanced survival. Contrary to our expectations, body mass was not significantly different between treatments at mating, birth of litter or weaning of young. Further, neither number nor size of young at weaning differed significantly between the two treatments. Survival to the next spring for adult females and juveniles was not significantly different between experimental and control treatments. Finally, annual fitness was not affected by the treatments. We concluded that females and their offspring were able compensate for the presence of ectoparasites, suggesting little or no fitness costs of parasites for females in the different colonies and during the years of our experiments.     

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.     

Endocrine profiles and reproductive output in European ground squirrels after unilateral ovariectomy

In the course of a long-term study on reproductive physiology and behaviour in female European ground squirrels (Spermophilus citellus) we investigated interactions among reproductive timing, maternal effort and hibernation. To gain information on the state of follicular development shortly before hibernation, histological investigations of ovaries were carried out. To collect the ovaries, unilateral ovariectomy was chosen as the least invasive method. The aim of this study was to monitor patterns of hormone secretion, behavioural parameters and reproductive output in females after unilateral ovariectomy, and compare them with intact individuals to assess potential effects of the treatment. Reproductive performance including duration of gestation and lactation, litter size, and offspring mass at natal emergence were similar in treated and untreated individuals. Before weaning, plasma estradiol levels were higher in the hemicastrated group than in intact females. Progesterone concentrations tended to be lower in unilaterally ovariectomized females during gestation and postlactation. In both groups we measured cortisol metabolites in fecal samples to estimate basal adrenal activity. Untreated individuals had significantly higher cortisol levels than hemicastrated females during lactation. In all other phases no significant differences in patterns of cortisol excretion were found between the two groups. Prehibernatory body-mass did not differ between treated and untreated females. These results indicate that unilateral ovariectomy had no significant negative effects on reproductive timing and output in female European ground squirrels. Compensatory activity of the remaining ovary does occur, concerning estradiol secretion even a kind of overcompensation was reported. These elevated estradiol levels in unilaterally ovariectomized females could play a role in limiting the degree of follicular compensation.     

Increasing returns in the life history of Columbian ground squirrels

1. We examined positive associations and trade-offs of maternal and reproductive traits in a population of Columbian ground squirrels, Spermophilus columbianus .
2. Structural size, body condition, mother's personal allocation to body mass during reproduction, and timing of littering were estimated for live-trapped reproductive females that were observed during an 8-year period, and were compared to litter mass, litter size, and average pup mass using path analyses.
3. Mothers exhibited age-structured traits that influenced reproductive patterns. Yearling mothers were significantly smaller, bred later, and had smaller litters than older females. Mothers that gained more body mass during reproduction and older mothers in good body condition that were structurally large had larger litters.
4. Early seasonal timing of littering was an important positive influence on successful reproduction by older mothers only in early breeding seasons and in years when conditions for reproduction were good for all females.
5. The number of offspring that survived to 1 year of age was most strongly associated with litter mass and litter size; date of breeding was of secondary influence, with earlier litters exhibiting greater success.
6. In general, mothers that gained the most in body mass during reproduction were concurrently more successful in weaning large litters (perhaps due to better quality of foraging habitat).
7. In addition to expected reproductive trade-offs, reproduction by Columbian ground squirrels exhibited positive associations of life-history traits that may reflect evolutionary increasing returns.     

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.     

Female red squirrels fit Williams' hypothesis of increasing reproductive effort with increasing age

Williams predicted that reproductive effort should increase as individuals age and their reproductive value declines. This simple prediction has proven difficult to test because conventional measures of energy expenditure on reproduction may not be a true reflection of reproductive effort. We investigated age-specific variation in female reproductive effort in a stable population of North American red squirrels where energy expenditure on reproduction is likely to reflect actual reproductive effort. We used seven measures of reproductive effort spanning conception to offspring weaning. We found that females completed growth by age 3 and that reproductive value decreased after this age likely because of reproductive and survival senescence. We therefore, predicted that reproductive effort would increase from age 3 onwards. The probability of breeding, litter mass at weaning, and likelihood of territory bequeathal were all lower for 1- and 2-year-old females than for females older than 3 years, the age at which growth is completed. That growing females are faced with additional energetic requirements might account for their lower allocation to reproduction as compared with older females. The probability of attempting a second reproduction within the same breeding season and the propensity to bequeath the territory to juveniles increased from 3 years of age onwards, indicating an increase in reproductive effort with age. We think this increase in reproductive effort is an adaptive response of females to declining reproductive values when ageing, thereby supporting Williams' prediction.     

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.     

Neonatal androgenization in ground squirrels: influence on sex differences in body mass and luteinizing hormone levels

Female squirrels were injected at birth with 50 or 1000 micrograms testosterone propionate (TP); control males and females were treated with oil vehicle. Squirrels were gonadectomized at 47 days of age. Body mass was recorded weekly and plasma luteinizing hormone (LH) was determined once monthly over the next year. Marked annual cycles in body mass were manifested by 30 out of 31 squirrels. Peak body mass and peak-to-trough differences were greater for control male and TP-female squirrels than for control female squirrels. Trough body weights did not differ among the groups. Luteinizing hormone was detectable in all male and most androgenized females but not in any control female squirrels during the first 4 mo after gonadectomy. Peak LH values were significantly greater for control male than for control female squirrels and were not influenced by neonatal androgenization in females. Testosterone propionate treatment also did not affect sex differences in timing of LH peaks or the total number of months in which LH was detectable. We conclude that testicular hormones secreted during the early postnatal period induce sex differences in the circannual pattern of weight change and some aspects of LH secretion. Complete masculinization, however, either requires more extensive action of gonadal hormones, perhaps both pre- and postnatally, or occurs through some androgen-independent mechanism.     

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.     

Age-specific variation in survival, reproductive success and offspring quality in red squirrels: evidence of senescence

Individual performance is expected to decrease with age because of senescence. We analyzed long-term data collected on a North American red squirrel population to assess the influence of age on body mass, survival and reproductive performance, and to study the effects of sex and of environmental conditions during early life on senescence patterns. Mass of males and females did not decrease at the end of life, possibly because body mass mostly reflects overall size in income breeders such as red squirrels. On the other hand, we found evidence of senescence in survival of both sexes and, to a lesser extent, in female reproductive traits. When compared to females, males had both higher survival and delayed decrease in survival, suggesting a weaker senescence in males. The offspring survival from weaning to one year of age also decreased with increasing mother age. This suggests that older females produce juveniles of lower quality, providing evidence of an intergenerational effect of mother's age on juveniles' fitness. Finally, our results indicate that variations in food conditions during early life influenced the reproductive tactics of females in the first years of their life, but not senescence patterns.     

Juvenile social play and yearling behavior and reproductive success in female Belding’s ground squirrels

Benefits of play behavior have been described for individuals during the juvenile period; however, it is less clear whether benefits of juvenile play commonly extend beyond the period of juvenile development. I evaluated possible associations between juvenile social play and yearling maternal territorial behavior and reproductive success in female Belding’s ground squirrels (Urocitellus beldingi). Higher rates of juvenile social play were associated with greater intensity of territorial aggressive and vigilant behavior during gestation but not later phases of the yearling reproductive cycle, suggesting that both juvenile play and later experience may influence maternal territorial behavior in yearling females. Juvenile social play was also a reliable predictor of whether a yearling female successfully weaned a litter, raising the possibility that juvenile play behavior influences yearling reproductive success via effects on maternal territorial behavior in female U. beldingi. Other factors were also related to reproductive success in yearling females. Body mass at the beginning of the yearling reproductive period was not correlated with juvenile social play but was associated with the likelihood of successfully weaning a litter. Yearling females whose mother was still alive were more likely to wean a litter than those whose mother was not alive. Mothers and yearling daughters tended to have maternal territories near each other, and mothers with a surviving yearling daughter expressed aggressive behavior at elevated rates and directed aggression toward intruders on the daughter’s territory. Overall, results of the study suggest that juvenile social play is among the variables that influence maternal territorial behavior and reproductive success in yearling female U. beldingi, and are consistent with the idea that juvenile play has benefits beyond the juvenile period for female U. beldingi.     

Short- and long-term effects of litter size manipulation in a small wild-derived rodent

Iteroparous organisms maximize their overall fitness by optimizing their reproductive effort over multiple reproductive events. Hence, changes in reproductive effort are expected to have both short- and long-term consequences on parents and their offspring. In laboratory rodents, manipulation of reproductive efforts during lactation has however revealed few short-term reproductive adjustments, suggesting that female laboratory rodents express maximal rather than optimal levels of reproductive investment as observed in semelparous organisms. Using a litter size manipulation (LSM) experiment in a small wild-derived rodent (the common vole; Microtus arvalis), we show that females altered their reproductive efforts in response to LSM, with females having higher metabolic rates and showing alternative body mass dynamics when rearing an enlarged rather than reduced litter. Those differences in female reproductive effort were nonetheless insufficient to fully match their pups’ energy demand, pups being lighter at weaning in enlarged litters. Interestingly, female reproductive effort changes had long-term consequences, with females that had previously reared an enlarged litter being lighter at the birth of their subsequent litter and producing lower quality pups. We discuss the significance of using wild-derived animals in studies of reproductive effort optimization.     

Reproductive activities of female albino rats treated with quassin, a bioactive triterpenoid from stem bark extract of Quassia amara

To evaluate the effect of quassin on female reproductive functions, 42 albino rats (35 females and 7 males) were used. The female albino rats were divided into seven groups of five rats each. Group I served as the control group and received distilled water while Groups II, III and IV rats were treatedorally with 0.1mg/kg, 1.0 mg/kg and 2.0 mg/kg body weight of quassin for 60 days respectively. Groups V, VI and VII rats were also treated orally with 0.1 mg/kg, 1.0mg/kg and 2.0 mg/kg body weight of quassin for 60 days but were left untreated for another 30 days, to serve as the recovery groups. At the end of each experimental period, blood samples were collected from each rat. Fertility study was done by cohabiting one untreated male with the five female rats in each group for 10 days. Quassin did not adversely affect the weight of the kidney, heart, liver and the body of the rats. However there was a significant decrease in the weight of the ovary and uterus in all the groups relative to the control. There was also a significant decrease in serum estrogen levels in quassin treated rats. The quassin treated rats had a significantly decreased mean litter number and weight. Histological studies show a disorganization and degeneration in the ovary while the uterus showed signs of vacuolation and disorganization. However, these effects were ameliorated after quassin was withdrawn from the rats. The results suggest that quassin has female anti-fertility properties, possibly acting via inhibition of estrogen secretion. Keyword: Quassin, Female rat, Reproduction, Estrogen.     

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.     

Effects of population density and female body mass on litter size in European roe deer at a continental scale

Roe deer (Capreolus capreolus) has a wide distributional range in Europe and inhabits a broad range of habitats and environmental conditions. Thus, populations of roe deer show substantial variation in demographic parameters. We aimed to determine whether body mass and population density—which influence the reproductive potential of roe deer at a local scale—affect their reproductive potential at a biogeographical scale as well. We reviewed the literature (covering years 1948–2015) on in utero litter size in roe deer from 59 locations in 14 countries in Europe. Across study sites, mean litter size varied from 1.0 to 2.4 embryo or corpora lutea per female, and population density ranged from 4.5 to 73.5 individuals/km². Mean body mass varied from 11.2 to 20.8 kg in subadult females and from 12.1 to 22.4 kg in adult females. Between 46° and 56° N, body mass of females did not show a significant trend of increase, whereas between 56° and 63° N, it increased with latitude (Bergmann’s rule). We used linear mixed-effects models (LMMs) to analyse the influence of body mass and population density (analysed separately) on litter size. Females with larger body mass produced larger litters, and this pattern was pronounced at both higher and lower latitudes. Higher population densities negatively affected litter size in a sample of all females. This macroecological analysis showed that factors influencing the reproductive potential of roe deer females at local scales produce similar effects at the biogeographical scale.     

Does gliding when pregnant select for larger females?

For terrestrial vertebrates, gliding imposes unique constraints on the interaction of body mass and structural size, particularly with reference to minimizing wing loading. Females of gliding animals experience increases in wing loading during pregnancy or gravidity, and selection may favour increased structural size to compensate for the added mass. We tested whether pregnant southern flying squirrels Glaucomys volans had similar wing loading as males, and whether females with lower wing loading bore heavier litters, than those with greater wing loading. Males had greater wing loading than females, regardless of the latter's reproductive state (males: 38.4±3.62 N m⁻², pregnant females: 30.7±4.21 N m⁻² and non-pregnant females: 26.8±5.13 N m⁻²). The slope of the linear relationship between planar surface area and body mass was similar between pregnant females and males, however ( F =0.383, P =0.322). Thus female flying squirrels may optimize their litter mass to minimize wing loading during pregnancy. Contrary to our prediction, females with greater wing loading had heavier litters than those with lower wing loading, which suggests reproductive output may be influenced by other ecological factors.     

Long-term effects of accelerated or delayed sexual maturation on reproductive output in wild female house mice (Mus musculus)

The effects of acceleration and delay of puberty in female house mice on survival and reproduction were tested using 6 experimental groups: (1) control females mated at the time of first oestrus, (2) females mated at weaning, (3) females treated with male urine starting at weaning and mated at first oestrus, (4) females housed in groups and mated at first oestrus, (5) females housed alone, treated with urine from grouped females and mated at first oestrus, and (6) females housed alone and mated at 68 days of age. Females caged with males at weaning or treated with male urine and mated at puberty had lower rates of survival to 180 days of age, but did not differ in rates of fertility from mice in the other four treatments. Those females that were housed with males from weaning or treated with male urine also had smaller total numbers of litters, fewer total young, and smaller average litter sizes than did females for which the age of mating was delayed, by grouping or treatment with urine from grouped females, or by being held until age 68 days before mating. Control females mated at first oestrus generally were intermediate or did not differ from the male treatments on these dependent variables. There were no differences in the average number of female young/litter across the 6 treatments. However, females that were delayed in age of first mating had significantly more male young/litter than did females that were accelerated in their sexual development or control females.(ABSTRACT TRUNCATED AT 250 WORDS)     

Reproductive effort and costs of reproduction in female European ground squirrels

Reproductive effort, factors affecting reproductive output and costs of reproduction were studied in primiparous yearling compared to multiparous older female European ground squirrels (Spermophilus citellus). Yearling females weaned smaller litters than older ones. Litter size increased with posthibernation body mass at the expense of slightly lighter young for yearling but not for older mothers. In older females, on the other hand, emergence body mass influenced offspring mass, whereas litter size was affected by oestrus date. High reproductive effort entailed reproductive costs in terms of reduced subsequent fecundity but not subsequent survival for both yearling and older females. The production of large litters and long duration of lactation delayed subsequent oestrus, which, in turn, correlated negatively with litter size. During the second half of lactation, oestradiol levels were significantly elevated, indicating the initiation of follicular maturation processes. Oestradiol levels during that time correlated negatively with current, but positively with subsequent litter size. We therefore assume that inhibitory effects of lactation on gonadal development may mediate the negative relationship between reproductive effort and subsequent reproductive timing in adults. This effect is absent in yearlings because they are reproducing for the first time. Reproductive output in yearlings was influenced by interactions between structural growth and puberty. Received: 22 March 1999 / Accepted: 7 June 1999     

Life history patterns in female moose (Alces alces): the relationship between age,body size,fecundity and environmental conditions

I examined the relationship between age, body size and fecundity in 833 female moose (Alces alces) from 14 populations in Sweden sampled during 1989–1992. Data on population density, food availability and climatic conditions were also collected for each population. Age and body mass were both significantly positively related to fecundity, measured as ovulation rate, among female moose. The relationship between the probability of ovulation and body mass was dependent on age with (1) a higher body mass needed in younger females for attaining a given fecundity, and (2) body mass having a stronger effect on fecundity in yearling (1.5 year) than in older (2.5 year) females. Thus, a 40 kg increase in yearling body mass resulted in a 42% increase in the probability of ovulation as compared to a 6% increase in older females. The lower reproductive effort per unit body mass, and the relatively stronger association between fecundity and body mass in young female moose compared to older ones, is likely to primarily represent a mechanism that trades off early maturation against further growth, indicating a higher cost of reproduction in young animals. In addition to age and body mass, population identity explained a significant amount of the individual variation in fecundity, showing that the relationship between body mass and fecundity was variable among populations. This variation was in turn related to the environment, in terms of climatic conditions forcing female moose living in relatively harsh/more seasonal climatic conditions to attain a 22% higher body mass to achive the same probability of multiple ovulation (twinning) as females living in climatically milder/less seasonal environments. The results suggests that the lower fecundity per unit body mass in female moose living in climatically harsh/more seasonal environments may be an adaptive response to lower rates of juvenile survival, compared to females experiencing relatively milder/less seasonal climatic conditions.     

Testing the reproductive and somatic trade‐off in female Columbian ground squirrels

Energetic trade‐offs in resource allocation form the basis of life‐history theory, which predicts that reproductive allocation in a given season should negatively affect future reproduction or individual survival. We examined how allocation of resources differed between successful and unsuccessful breeding female Columbian ground squirrels to discern any effects of resource allocation on reproductive and somatic efforts. We compared the survival rates, subsequent reprodction, and mass gain of successful breeders (females that successfully weaned young) and unsuccessful breeders (females that failed to give birth or wean young) and investigated “carryover” effects to the next year. Starting capital was an important factor influencing whether successful reproduction was initiated or not, as females with the lowest spring emergence masses did not give birth to a litter in that year. Females that were successful and unsuccessful at breeding in one year, however, were equally likely to be successful breeders in the next year and at very similar litter sizes. Although successful and unsuccessful breeding females showed no difference in over winter survival, females that failed to wean a litter gained additional mass during the season when they failed. The next year, those females had increased energy “capital” in the spring, leading to larger litter sizes. Columbian ground squirrels appear to act as income breeders that also rely on stored capital to increase their propensity for future reproduction. Failed breeders in one year “prepare” for future reproduction by accumulating additional mass, which is “carried over” to the subsequent reproductive season.