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     

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.     

Reproductive resilience to food shortage in a small heterothermic primate

The massive energetic costs entailed by reproduction in most mammalian females may increase the vulnerability of reproductive success to food shortage. Unexpected events of unfavorable climatic conditions are expected to rise in frequency and intensity as climate changes. The extent to which physiological flexibility allows organisms to maintain reproductive output constant despite energetic bottlenecks has been poorly investigated. In mammals, reproductive resilience is predicted to be maximal during early stages of reproduction, due to the moderate energetic costs of ovulation and gestation relative to lactation. We experimentally tested the consequences of chronic-moderate and short-acute food shortages on the reproductive output of a small seasonally breeding primate, the grey mouse lemur (Microcebus murinus) under thermo-neutral conditions. These two food treatments were respectively designed to simulate the energetic constraints imposed by a lean year (40% caloric restriction over eight months) or by a sudden, severe climatic event occurring shortly before reproduction (80% caloric restriction over a month). Grey mouse lemurs evolved under the harsh, unpredictable climate of the dry forest of Madagascar and should thus display great potential for physiological adjustments to energetic bottlenecks. We assessed the resilience of the early stages of reproduction (mating success, fertility, and gestation) to these contrasted food treatments, and on the later stages (lactation and offspring growth) in response to the chronic food shortage only. Food deprived mouse lemurs managed to maintain constant most reproductive parameters, including oestrus timing, estrogenization level at oestrus, mating success, litter size, and litter mass as well as their overall number of surviving offspring at weaning. However, offspring growth was delayed in food restricted mothers. These results suggest that heterothermic, fattening-prone mammals display important reproductive resilience to energetic bottlenecks. More generally, species living in variable and unpredictable habitats may have evolved a flexible reproductive physiology that helps buffer environmental fluctuations.     

Reproductive performance in female common hamsters

We monitored individual reproductive timing and output in a common hamster population in Vienna over a 3-year period. Animals were live-trapped, weighed, individually marked, and reproductive status was determined at capture. Costs of reproduction were investigated by measuring body condition shortly before hibernation and overwinter survival. Our results indicated that early emerging females had more litters and weaned more offspring per season. Body mass throughout the active season did not differ significantly between females with high and low reproductive output. High reproductive output seemed to affect the duration of the active season. Successful females had a longer postreproduction period before hibernation than less successful ones, probably serving to balance the costs of reproductive effort by extended preparation for hibernation. Also, females that had emerged early in spring and had high reproductive success were more likely to survive the subsequent winter. Hence, we found female common hamsters to vary strongly in maternal investment capacity and to tailor reproductive strategies accordingly.     

Parameters of Mating Effort and Success in Male European Ground Squirrels,Spermophilus citellus

Behavioural and physiological changes during the reproductive period were documented in male European ground squirrels including: initiated conflict, scent marking, locomotion and range size, as well as testis size, scrotal pigmentation, testosterone levels and changes in body mass. Spring emergence dates in adult males varied but preceded those of females. Gonadal maturation was complete in adult males at emergence. Most yearlings did not mature and emerged with females. After a quiescent premating phase, male range, range overlap, conflict rates and locomotion increased at female emergence. The synchrony of females, their small ranges and short latencies to ovulation produced a highly male-biased operational sex ratio (OSR) and potential for scramble competition polygyny. In contrast to other ground squirrel species, each female apparently mated with only a single male. Scramble competition did appear to occur, in that males tried to contact as many females as often as possible before copulation. Females placed high temporal prerequisites on males. Costs of scrambling were documented by body mass loss during mating, which covaried with the number of ‘acquired’ females. Testicular regression occurred earlier, and fattening, moult and onset of next hibernation were delayed in successful males. The determinant factor in these costs was found to be mass loss intensity during the main mating period. Surprisingly, this factor was best related to feeding changes and not to locomotion or aggression. Time budget constraints perhaps imposed by the female, with compounding influences of exercise and stress effects of conflict, then negatively affected ensuing seasonal activities. The individual changes may adversely affect individual survivability and/or future reproductive success.     

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.     

Sex and the Single Vole: Effects of Social Grouping on Prairie Vole Reproductive Success

Prairie voles (Microtus ochrogaster) have a mating system that is primarily monogamous with paired males and females together defending breeding space against intruders of either sex. Breeding success may be affected when other adults intrude on the territorial space of pairs. We conducted an experiment to determine the impact of additional members of either sex on reproductive success of pairs. In laboratory arenas, we formed pairs (1F:1M) and two kinds of triad (2F:1M, 1F:2M). Females in pairs had the highest conception rates, litter sizes and survival of litters. Females in 1F:2M groups had slightly reduced litter sizes and reduced numbers of weanlings, and some females had litters sired by both males. Females in 2F:1M groups had low conception rates and the smallest litters, and >35% of their litters suffered infanticide; in no case did both females become pregnant. Throughout the trials, individuals of the sex doubly represented in triads were more likely to die than were individuals of the sex singly represented. We conclude that there may be fitness costs associated with the presence of unrelated supernumerary adults during gestation and lactation.     

Hormonal manipulation of offspring number: maternal effort and reproductive costs

We used exogenous gonadotropin hormones to physiologically enlarge litter size in the bank vole (Clethrionomys glareolus). This method allowed the study design to include possible production costs of reproduction and a trade-off between offspring number and body size at birth. Furthermore, progeny rearing and survival and postpartum survival of the females took place in outdoor enclosures to capture salient naturalistic effects that might be present during the fall and early winter. The aim of the study was to assess the effects of the manipulation on the growth and survival of the offspring and on the reproductive effort, survival, and future fecundity of the mothers. Mean offspring body size was smaller in enlarged litters compared to control litters at weaning, but the differences disappeared by the winter. Differences in litter sizes disappeared before weaning age due to higher mortality in enlarged litters. In addition to the effects of the litter size, offspring performance was probably also influenced by the ability of the mother to support the litter. Experimental females had higher reproductive effort at birth, and they also tended to have higher mortality during nursing. Combined effects of high reproductive effort at birth and high investment in nursing the litter entailed costs for the experimental females in terms of decreased probability of producing a second litter and a decreased body mass gain. Thus, enlarged litter size had both survival and fecundity costs for the mothers. Our results suggest that the evolution of litter size and reproductive effort is determined by reproductive costs for the mothers as well as by a trade-off between offspring number and quality.     

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.     

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.     

Age-specific resource investment strategies: evidence from female Richardson's ground squirrels (Spermophilus richardsonii)

To avoid a possible cost to their future survival and/or reproduction, individuals must balance their somatic and reproductive investments. The van Noordwijk and De Jong model of resource investment predicts that investments into reproduction and soma can vary among individuals of a population based on the variation in the total amount of energy that individuals acquire. With principal components analysis (PCA), we created two axes of life history for female Richardson's ground squirrels Spermophilus richardsonii : an index of total energy investment (PC1) and an index of investment tactic (PC2). Using these indices, we examined patterns of resource allocation to reproductive and somatic investments. Because yearling female Richardson's ground squirrels complete growth to adult size during pregnancy and early lactation, their somatic needs exceed those of older, fully grown females. Therefore, we predicted that yearlings would show more evidence of a tradeoff between reproductive and somatic investments compared with older females. Both yearling and older females invested four to five times more mass into their litters than into their own body mass. With increasing total investment, yearling females increased investment in both reproduction and themselves, whereas older females invested relatively more in reproduction than themselves. Regardless of age, females that emerged heavier from hibernation invested fewer resources into themselves and more into their litters. Variation in total energy investment and investment tactic indices was similar for yearling and older females. Contrary to our prediction, however, yearling females showed positive associations between reproductive and somatic investments, whereas older females exhibited showed no significant association between reproductive and somatic investments.     

Variation in reproductive investment among and within populations of the scorpion Centruroides vittatus

Although information concerning variation among and within populations is essential to understanding an organism's life history, little is known of such variation in any species of scorpion. We show that reproductive investment by the scorpion Centruroides vittatus varied among three Texas populations during one reproductive season. Females from the Kickapoo population produced smaller offspring and larger litters than females from the Independence Creek or Decatur populations; this pattern remained when adjusting for among population variation in either female mass or total litter mass. Relative clutch mass (RCM) and within-litter variability in offspring mass (V*) did not differ among populations. Among-population variation may result from genetic differences or from phenotypically plastic responses to differing environments. Within populations, the interrelationships among reproductive variables were similar for Decatur and Independence Creek: females investing more in reproduction (measured by total litter mass, TLM) produced larger litters and larger offspring, and V* decreased with increased mean offspring mass (and with decreased litter size at Decatur). At Kickapoo, larger females produced larger litters and had larger TLM; females investing more in reproduction produced larger litters but not larger offspring. Within litter variability in offspring mass was not correlated with any reproductive variables in this latter population. These patterns may be explained by the fractional clutch hypothesis, the inability of females precisely to control investment among offspring or morphological constraints on reproduction.     

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.     

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.     

南滑蜥繁殖行为和繁殖策略的研究

南滑蜥Scincella reevesii 系温暖气候区卵胎生蜥蜴,每年繁殖1次,繁殖季节为5月下旬到6月上旬.平均每只繁殖雌蜥产仔6(3～9)条.雌蜥分娩持续时间为30～ 60 min,仔蜥在产出时包被卵膜.南滑蜥雌体采取高投入的繁殖策略,平均繁殖投入为0.359,平均窝仔投人为0.247.相关性回归分析表明,窝仔数、窝仔重与雌蜥体长呈正相关关系,幼仔重与雌蜥体长不相关;窝仔数与雌蜥的繁殖投入及窝仔投入均成正相关,与分娩时的能耗投入不相关,与幼仔投入成显著负相关.     

Reproductive investiment in the scorpion Centruroides vittatus

Among invertebrates, scorpions possess a relatively unique set of reproductive traits. The interrelationships of these traits may have important implications for life history theory, yet there have been few studies of these traits in scorpions. Our data indicate that larger female Centruroides vittatus produce more offspring and have a higher total litter mass than smaller females. There was, however, no significant relationship between offspring size and female or litter size. Mean offspring mass increased with increases in total litter mass and within litter variation in offspring size (coefficients of variation) decreased with increasing total litter mass. These results suggest that large female scorpions with a larger investment in reproduction produced more offspring that were more uniform in size, but not significantly larger, than small females with less investment. The fractional clutch principle and physiological and functional constraints on size and number of offspring are suggested as possible explanations for the relationships we found among offspring size, variation in offspring size and total investment in offspring in C. vittatus.     

Reproductive capacity and dominant lethal mutations in female guinea-pigs and Djungarian hamsters following X-rays or chemical mutagens.

The reproductive capacity and induction of dominant lethal mutations in adult female guinea-pigs and Djungarian hamsters were tested following treatment with 400 rad X-rays, 1.6 mg/kg triethylenemelamine (TEM) or 75 mg/kg isopropylmethanesulphonate (IPMS). A fairly high level of dominant lethals were observed in female guinea-pigs mated at the first oestrus after irradiation (23.4 +/- 6.4%) with a lower yield at 3 months (9.6 +/- 8.2%). Neither of the chemicals caused any significant induction of dominant lethals at either mating time. In the reproductive capacity experiments, the mean litter size of irradiated female guinea-pigs was reduced for about 12 months and this was especially marked in the first 6 months following treatment. Neither of the chemicals caused any significant differences in early litter sizes but there was a noticeable reduction in the litter sizes of TEM-treated females in the 18--24 month interval. With Djungarian hamsters a marked effect of X-rays on reproductive capacity was apparent. After 400 rad a smaller proportion of irradiated females littered in the first 25-day interval than after the other treatments, and no irradiated females produced more than one litter. Neither of the chemicals caused such a drastic reduction in fertility but TEM-treated females produced fewer litters and became sterile at an earlier age than control or IPMS-treated females. With IPMS, the number of litters produced was similar to the controls. Both chemicals caused a significant reduction in litter-size but further work is needed to establish whether this was due to induction of dominant lethals. No translocations were observed in the sons of treated female guinea-pigs or hamsters, but the numbers of animals studied were too small for any conclusions to be drawn.     

Maternal investment during pregnancy in wild meerkats

Maternal investment in offspring development is a major determinant of the survival and future reproductive success of both the mother and her young. Mothers might therefore be expected to adjust their investment according to ecological conditions in order to maximise their lifetime fitness. In cooperatively breeding species, where helpers assist breeders with offspring care, the size of the group may also influence maternal investment strategies because the costs of reproduction are shared between breeders and helpers. Here, we use longitudinal records of body mass and life history traits from a wild population of meerkats (Suricata suricatta) to explore the pattern of growth in pregnant females and investigate how the rate of growth varies with characteristics of the litter, environmental conditions, maternal traits and group size. Gestational growth was slight during the first half of pregnancy but was marked and linear from the midpoint of gestation until birth. The rate of gestational growth in the second half of pregnancy increased with litter size, maternal age and body mass, and was higher for litters conceived during the peak of the breeding season when it is hot and wet. Gestational growth rate was lower in larger groups, especially when litter size was small. These results suggest that there are ecological and physiological constraints on gestational growth in meerkats, and that females may also be able to strategically adjust their prenatal investment in offspring according to the likely fitness costs and benefits of a particular breeding attempt. Mothers in larger groups may benefit from reducing their investment because having more helpers might allow them to lower reproductive costs without decreasing breeding success.     

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.     

Vaginal oestrus during the reproductive and non-reproductive period in European ground squirrels

Most hibernating animals are thought to be monoestrous because reproductive activity is subject to strong time constraints. In previous studies, female European ground squirrels (Spermophilus citellus) turned out to have elevated oestradiol levels during late lactation and after weaning, indicating ovarian activity during summer. Therefore, we monitored vaginal cytology, endocrine changes, and vulval development in semi-free-living female European ground squirrels throughout one active season. Vaginal oestrus, defined by the predominance of cornified cells in smear samples, was found during the mating period shortly after vernal emergence. This phase was followed by metoestrus, characterised by the appearance of leukocytes, and a subsequent anoestrous phase. During weaning or postlactation, a second vaginal oestrus was documented in all experimental females, again followed by a metoestrous and an anoestrous phase lasting until hibernation. In line with the second vaginal oestrus, plasma oestradiol concentrations peaked during postlactation. Progesterone levels were elevated from gestation to postlactation, and titres were marginally higher during vaginal oestrus in summer than in spring. Vulval swelling was more pronounced during the first than the second vaginal oestrus. The second oestrous cycle was non-reproductive, as males were sexually inactive with regressed testes during summer. We assume that the second oestrous cycle and the accompanying endocrine changes have beneficial effects on prehibernatory fattening and reproductive performance in the subsequent season. This might allow females to become oestrous immediately after emergence from hibernation in spring.