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

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

Litter reductions reveal a trade-off between offspring size and number in brown bears

Experimental manipulations have revealed positive effects of litter reduction on offspring mass in small mammals, but little is known about this trade-off in large mammals. We examined the determinants of natural litter size variation and quantified the effects of litter size, maternal characteristics, and litter composition on yearling mass using 24?years of data on marked brown bears (Ursus arctos) in Sweden. Infanticide by adult males is a major cause of cub-of-the-year mortality, leading to litter size reductions. Litter size (n?=?265) at den emergence ranged from one to four cubs (average, 2.7) and increased with maternal age. Litter size, however, appeared independent of maternal size, population density, interlitter interval, study area, or previous litter sex ratio. Yearling body mass increased with maternal body size but was independent of litter sex ratio. Litter size and yearling mass were negatively correlated, mostly because singletons were about 30?% heavier than yearlings from litters of two to four cubs. In reduced litters, survivors were on average 8?% heavier as yearlings than individuals from intact litters, suggesting that sibling competition reduces growth. Trade-offs between litter size and yearling mass in bears appear similar in magnitude to those found in small mammals.     

Offspring sex ratio variation in the bridled nailtail wallaby, Onychogalea fraenata

The bridled nailtail wallaby is a sexually size dimorphic, promiscuous, solitary macropod. Sex ratios of pouch young were studied at two sites over 3 years, beginning with 14 months of severe drought. Females that were in better condition were more likely to have sons, and condition was dependent on body size. Females at one site were heavier, were consequently in better condition, and produced more sons than females at the other site. Females that declined in condition had more daughters during the most severe part of the drought than females that maintained condition, but endoparasite infection did not affect the pouch young sex ratio. Age also appeared to affect sex ratio adjustment, because weight was strongly influenced by age. Sex ratio bias was not caused by early offspring mortality, but occurred at conception. Mothers did not appear to bias energy expenditure on sons or daughters; males and females did not differ in condition at the end of pouch life. Pouch young sex ratio variation was most consistent with the Trivers-Willard hypothesis, but could also have been influenced by local resource competition, since sons dispersed further than daughters. Offspring condition was related to survival, and was correlated with maternal condition. Received: 14 April 1998 / Accepted after revision: 10 November 1998     

Why do female Belding's ground squirrels disperse away from food resources?

We examined the effects of food provisioning on the natal dispersal behavior of Belding's ground squirrels (Spermophilus beldingi). We provided extra food to adult and yearling females in their maternal territories during pregnancy and lactation, and to offspring of these females in their natal areas for 6 weeks after weaning. We used unprovisioned young of unprovisioned mothers as controls. Provisioning influenced the probability of dispersal from the natal area by female but not male S. beldingi. All surviving male S.␣beldingi dispersed by 55 weeks of age, regardless of whether they and their mothers received extra food. By contrast, we observed a significant trend, beginning 3 weeks after weaning and continuing through the yearling year, for a greater proportion of provisioned than control female S. beldingi to emigrate from the natal area. Competition for food did not appear to influence natal dispersal of females. However, overall population density, density of females weaning litters, and rates of aggression and vigilance among these females, were higher in provisioned than control areas, suggesting that competition for non-food resources was unusually intense in provisioned areas. We propose that juvenile female, but not juvenile male, S. beldingi may emigrate from the natal site to increase access to areas with low densities of conspecifics. Together with findings of earlier workers, our results suggest that spatial and temporal distributions of environmental resources are important influences on the dispersal behavior of female ground squirrels. Received: 28 February 1996 / Accepted after revision: 16 October 1996     

Inbreeding-based bias in parental responsiveness to litters of oldfield mice

This study investigated variation in parental behavior in oldfield mice (Peromyscus polionotus). Specifically, I examined the possibility that parental behavior and probability of litter loss may differ depending on whether parents are related to one another and, therefore, whether offspring are inbred. When parents are related, they share more genes with their offspring and so may be predicted to invest more in inbred offspring if these offspring do not inherently have a greater risk of juvenile mortality. Survival of inbred pups did not differ from survival of outbred pups, and females tended to exhibit a preference, in the form of enhanced parental care, for inbred litters. Males did not display any consistent preferences. Inbred litters were left unattended more often during the immediate post-partum period than were outbred litters, but this inconsistent finding appeared to have a smaller effect than the overall improved maternal care provided to inbred litters. Females thus appeared to modulate their behavior to a greater degree than did males depending on characteristics of the litter. Patterns of litter loss indicated that complete death of litters was independent of litter inbreeding, whereas loss of selected pups in a litter occurred significantly more often when litters were inbred. Complete and partial loss of litters may be functionally different behaviors that are triggered by different cues. Received: 20 September 1996 / Accepted after revision: 28 April 1997     

The association of female kin in the arctic ground squirrel Spermophilus parryii

Summary The association of kin in arctic ground squirrels (Spermophilus parryii) was studied near Haines Junction, Yukon Territory, during the summers of 1977, 1978, and 1979. Males dispersed in this population, thus only females were likely to live near relatives.Close female kin (sisters, mothers/daughters) had greater overlap of home ranges and interacted more amicably and less agonistically than did less closely related females. Closely related females clumped their young at emergence, whereas more distantly related females did not; there was little indication of clumping of pre-emerged young. The overlap of home ranges of distant relatives (known genetic relatives that had not associated in a natal burrow) was intermediate to that of close relatives and non-relatives. The types of interactions between distant relatives were more similar to those between non-relatives than between close relatives.I conclude that female arctic ground squirrels exhibit nepotism. Females may benefit from associations with relatives by sharing watching for predators once juveniles become conspicuous. Male arctic ground squirrels commit infanticide and several females may be more effective at protecting their young from infanticidal males than females living alone. I suggest that clumping of young by close relatives may provide a mechanism allowing distantly related females to identify each other.Address for offprint requests     

Components of lifetime reproductive success in communally and solitarily nursing house mice — a laboratory study

Under laboratory conditions, communal nursing among familiar and closely related female house mice (Mus domesticus) improved lifetime reproductive success compared to females rearing litters alone or females living with a previously unfamiliar, unrelated partner (reproductive success was measured within an experimental lifespan of 6 months, standardized as 120 days after mating at the age of about 2 months). An analysis of the contribution of three multiplicatively combined components to variation in reproductive success among breeding females revealed that, in all three social groups, survival of young until weaning contributed most to differences in lifetime reproduction (46–64% of the total variance). Females living with a sister had a significantly higher probability of reproducing successfully than females in the other groups, and also reared significantly more litters communally than females sharing nests with an unrelated partner. Weaning probabilities of young were highest in litters cared for by sisters and lowest in nests of unrelated females. Young were found dead either directly after birth (within the first 2 days of lactation) or after they had been cared for and nursed for at least 1 day. The loss of an entire litter typically occurred directly after birth. In monogamous females rearing litters alone the death of almost all young coincided with such early entire-litter mortality. In polygynous groups, however, offspring died at an older age and more litters suffered the loss of some young. Still, rearing young with a sister improved survival directly after birth and fewer litters were lost entirely in comparison with females in the other groups. In polygynous groups, pregnant females were observed to kill some of their partner's dependent young shortly before they gave birth themselves. As a consequence, individual young had reduced survival when they were firstborn in a communal nest (another litter was born within 16 days). Analyzed over a lifetime, communal care among familiar and closely related female house mice seems to be an adaptation to maximize the survival of offspring until weaning.     

Sex ratio manipulation and decreased growth of male offspring of undernourished golden hamsters (Mesocricetus auratus)

Summary The theory that female mammals in poor condition may increase individual fitness by skewing the sex ratio of their offspring toward daughters and by investing more resources in daughters than in sons was tested in hamsters. Newly mated experimental females were food-restricted during pregnancy and lactation (RR) or during lactation only (AR). Controls received food ad libitum. Maternal body weights were assessed daily from mating to 25 days postpartum. Litter survival (% litters with at least one pup surviving on any day), litter size, offspring sex ratios (=% males), and pup weights were monitored daily from birth (Day 1) to Day 25. All control and AR dams gave birth 16 days after mating. Gestation was extended by 1–3 days for 35.4% of RR dams. RR dams weighed significantly less at parturition than controls and AR females. During lactation, AR females showed the greatest weight loss and control females the least. AR weight loss exceeded that of RR females, possibly because the former maintained larger litters. Survival was highest for controls, intermediate for AR, and lowest for RR litters. Mean sex ratio at birth was significantly less for RR (40.7%) than for control (49.6%) and AR (48.8%) litters. RR sex ratio did not change significantly postnatally. Sex ratios of control and AR litters never differed statistically from 50%. Control male pups were significantly heavier than their sisters throughout the experiment. No significant gender differences were observed for AR pup weights after Day 2 postpartum. RR female offspring weighed more than their brothers throughout the experiment, and this difference was statistically significant immediately prior to the time that pups began to feed independently (Days 14–17). RR female pup weights were similar to, and sometimes significantly greater than, weights of control daughters during the period of postnatal maternal investment. Control males were always heavier than males from the other treatments. Patterns of weight gain by AR and RR males varied with age. We conclude that underfed female hamsters are able to adjust the sex ratio of offspring prenatally and parental investment postnatally to favor daughters.     

Emigration mechanisms in feral house mice – a laboratory investigation of the influence of social structure,population density,and aggression

Emigration in small mammals may be strongly related to social factors, but direct observations of emigrants are rare. Feral house mice (Mus domesticus) were studied using a population cage system that allowed continuous observation of individually marked animals. Mice that left their natal cage and took up residence in cages that could only be reached by crossing a water barrier were defined as emigrants. Six pairs of house mice with their litters were placed in the system, and data on aggressive interactions, body weight, reproduction, mortality and emigration were collected daily. Both sexes emigrated, but males did so twice as often as females. Population density was not correlated with the frequency of aggression, and had no effect on the weight of emigrating individuals. Male emigrants suffered more aggression before emigration than their non-emigrant brothers of the same age; they were aggressively driven out by other males, predominantly by the father. Female emigration depended on the female’s chances of reproduction. The probability of a female reproducing decreased with increasing birth order. Females born in a late litter, who therefore had only a low chance of reproduction, dispersed earlier than those of early litters. Resident males were reproductively suppressed. Male offspring had two different strategies for attaining top rank. They could develop rapidly and reach sexual maturity early on, but face competition with the father, risking being forced to emigrate. Alternatively, they could develop slowly, stay within their family and wait for a chance to take over the dominant position. It is concluded that emigration in male and female feral house mice is caused by intrasexual competition. Received: 13 July 1995/Accepted after revision: 8 June 1996     

The use of incubation behavior to adjust avian reproductive costs after egg laying

Reproduction in birds requires the input of time and energy during discrete breeding phases leading to investment trade-offs between laying date, clutch size, body mass, and incubation constancy. We investigated costs during incubation by experimentally enlarging 25 clutches of white-tailed ptarmigan Lagopus leucurus. The experiment was conducted in 2 years, one with harsh weather that forced a natural delay in reproduction. When forced to delay egg-laying, females began incubation with poorer body condition and foraged more during incubation. Rates of mass loss during incubation were not affected by clutch enlargement, and did not differ between harsh and benign years; however, females that were heavier at the start of incubation lost more mass than lighter females. Clutch-enlarged females had reduced nest attendance compared to control birds in both years and incubation periods increased by up to 2 days relative to controls. In the harsh year, there was a trend for clutch-enlarged females to have lower nest success, but there was no effect on overwinter survival. Different behavioral responses by females in the 2 years showed that incubation costs may depend on other factors such as female quality, food supply, or weather conditions. Incubation is a dynamic period during which birds may adjust energy balances by varying body condition and food intake. Received: 28 March 2000 / Received in revised form: 18 August 2000 / Accepted: 2 July 2000     

Maternal care in house mice

Summary This paper analyzes the flexibility of maternal care in wild house mice (Mus domesticus) under different reproductive conditions in the laboratory. All maternal activities were both qualitatively and quantitatively analyzed over a period of 28 days after birth of a litter. The standard behavior of a lactating house mouse with 7–8 young can be described as follows: During days 1–16 the offspring fully depend on the mother for nutrition. Due to rapid growth of the litter, the energetic demands of lactation reach a peak for the female during days 13–16. During days 17–22, the weaning period, the young begin to eat solid food. This period is characterized by behaviors that indicate different interests of the mother and offspring, and thus the existence of a parent-offspring conflict sensu Trivers (1974). Resting alone and remaining far from the litter indicate the female's interest in avoiding the offspring's demands, which are expressed in frequent attempts to initiate sucking. There is no aggression towards the young during weaning. House mice are weaned at 23 days. The relationship between mother and young appears free of conflict after weaning. Nursing is replaced by resting with body contact, but the offspring do not try to suck. The following results suggest that during the weaning period the offspring do not get more milk than corresponds to the maternal optimum—despite their frequent sucking attempts:(a) When the mother is simultaneously lactating and pregnant, offspring are smaller at weaning than under standard conditions. (b) Small litters are weaned earlier than large ones. Despite a longer nursing period, offspring from large litters are lighter at weaning than those from small ones. (c) Under high energy demand, as after postpartum mating and with large litters, females wean their young at a body weight which corresponds to the earliest physiologically possible state of independence.Parity of the female has no effect on maternal activities, nor has the presence of the father. In the latter case, however, offspring are less often left alone and unprotected.Females seem to adjust their investment according to the body weight of the progeny by delaying or advancing the date of weaning (Table 2). This behavior allows the production of the largest possible number of offspring that can be raised to a minimal physiological threshold corresponding to a body weight of approximately 9 g. Such flexibility in parental care may enhance maternal fitness under different and unpredictable environmental conditions.     

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

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

Time constraints and multiple choice criteria in the sampling behaviour and mate choice of the fiddler crab,Uca annulipes

Active female sampling occurs in the fiddler crab Uca annulipes. Females sample the burrows of several males before remaining to mate in the burrow of the chosen partner. Females time larval release to coincide with the following nocturnal spring tide and must therefore leave sufficient time for embryonic development after mating. Here we show how this temporal constraint on search time affects female choosiness. We found that, at the start of the sampling period (when time constraints are minimal), females selectively sample the larger males in the population. Towards the end of the sampling period (when the temporal constraints increase the costs of sampling), females are less selective. Furthermore, we suggest that the number of males sampled (and other indices of ‘‘sampling effort’’) may not be reliable indicators of female choosiness and may not reflect the strength of female mating preferences under certain conditions. Burrow quality also emerged as an important criterion in final mate choice. Burrow structure potentially influences reproductive success, and mate acceptance based on burrow structure appears to involve a relatively invariant threshold criterion. Since there is no relationship between male size and burrow quality, females are using at least two independent criteria when choosing potential mates. We envisage mate choice as a two-stage process. First, females select which males to sample based on male size. They then decide whether or not to mate with a male based on burrow features. This sampling process explains how two unrelated variables can both predict male mating success. Received: 23 March 1995/Accepted after revision: 14 January 1996     

Maternal investment, sex-differential prospects, and the sex ratio in wild house mice

In a population of first-generation offspring from wild-caught house mice (Mus musculus domesticus), previous evidence suggested that male fitness is more strongly affected by an increase in body weight than female fitness. This paper shows that in these mice the young are weaned at heavier weights the smaller the litter and the better the maternal body condition. These effects persisted into adulthood and were less pronounced in female young. However, contrary to expectation from conventional sex ratio theory, maternal condition and litter size had no detectable effect on sex ratios. Also, litter size did not affect sex ratios in two populations of laboratory-kept, wild-caught western (M. m. domesticus) and eastern house mice (M. m. musculus). Wild house mice, therefore, appear not to adaptively manipulate the sex ratio of offspring. It is argued that this absence of sex ratio trends might not be maladaptive, but rather that models currently used to predict sex ratio trends in rodents may not be valid. Received: 13 March 1997 / Accepted after revision: 9 August 1997     

Physical variability among nest foundresses in the polygynous social wasp,Polistes annularis

Summary An individual's rank in a dominance hierarchy is often based on size or weight, especially in insects. Foundresses of the social wasp, Polistes annularis, vary greatly in size as measured by wing length, dry weight, fat weight, and residue weight after fat has been extracted. Females that emerged from the same nest are much more similar in size to each other than they are to females that emerged from other nests. Within nests however queens are usually larger than their subordinates. Queens that emerged from one nest may be smaller than subordinates that emerged from another nest. We found no evidence of a group of females that are forced into being subordinates because of inadequate feeding as larvae. Females are also probably not attempting to begin new nests with females as different in size as possible from themselves since means and variances of winglength of females on new nests do not differ from those of all females emerging from that natal nest.     

Philopatry and mother-daughter associations in bushy-tailed woodrats: space use and reproductive success

Social groups in mammals are often based on overlapping generations of philopatric females, yet few researchers have examined fitness consequences of associations between females and their daughters. We examined survival and reproductive consequences of sharing a natal site with one's mother or daughter in wild bushy-tailed woodrats (Neotoma cinerea). Bushy-tailed woodrats reside on discrete rocky outcrops, and females tend to spend their entire reproductive life on their natal outcrop. Mothers and daughters that co-existed on the natal outcrop were closely associated in space. Juvenile females were more likely to survive their first winter to become breeding yearlings at their natal site if their mother was also present at the natal site during the winter. Juveniles survived equally well with or without their mother where densities of adult females were experimentally reduced, suggesting that females enhance their daughter's survival by facilitating their access to limited resources. Yearling females exhibited no reproductive costs from association with their mothers at the natal site during the breeding season; in fact, yearlings that bred at the natal site simultaneously with their adult mothers tended (P = 0.07) to experience fewer reproductive failures than did those that bred in the absence of their mothers. Mothers did not experience any detectable costs when sharing a natal site with a daughter. We conclude that bushy-tailed woodrats exhibit stable, fitness-enhancing associations among mothers and their philopatric daughters.     

Pup production,sex ratios,and survivorship in African wild dogs, <Emphasis Type="Italic">Lycaon pictus</Emphasis>

The local resource enhancement (LRE) model predicts that in cooperatively breeding species, sex ratios will be biased in favor of the more helpful sex. In this study, we assess the assumptions underlying the LRE model in a population of cooperatively breeding wild dogs (Lycaon pictus) in Northern Botswana monitored over a 15-year period. In this population, litter size and pup survival to 1 year are strongly affected by pack size and the breeding female’s age, but adult males have a stronger and more linear effect on females’ reproductive performance than do adult females. This asymmetry in the benefits derived from male and female helpers is reflected in male-biased sex ratios in litters at the time pups emerge from the den. Sex ratio biases are most pronounced in the litters of the youngest mothers who live in significantly smaller packs than older females. The presence of potential rivals for the dominant female’s position depresses pup production at the time of emergence, suggesting that competition among females for breeding positions may also contribute to the selective forces affecting birth sex ratios.     

Home range shifts by breeding female Townsend's voles (Microtus townsendii ): a test of the territory bequeathal hypothesis

Numerous accounts of partial or complete breeding dispersal by adult females reported in the literature on microtines have been interpreted as maternal bequeathal. However, for most species, no causal relationship between a female's movements between successive litters and the settlement of her offspring in the natal range has been demonstrated. I tested predictions from the bequeathal hypothesis using data from an intensive study of Townsend's voles where genetical relationship had been determined by marking most pups in the maternal nest before weaning. After correcting statistically for temporal changes in the distance moved between successive litters I rejected the following predictions: (1) that females with one or several daughters of reproductive age should be more likely to disperse or disperse farther between successive births than females without such daughters; (2) that mothers should be more likely to bequeath their range to their daughters in spring when the latter would gain most from having ready access to a breeding range; (3) that daughters benefit from their mother abandoning the natal range through an increase in their probability of recruitment; and (4) that daughters actually use the home range vacated by their mother. I critically reassess the empirical studies quoted as evidence that breeding dispersal is a form of parental investment in microtines and other mammals. Most empirical studies cited in support of the bequeathal hypothesis often only contain anecdotal reports of movements by breeding females or do not mention it as one of its possible adaptive functions. Some studies contain evidence only consistent with the bequeathal hypothesis whereas others are incompatible with this explanation. Documented evidence of bequeathal comes exclusively from species which rely on a semi-permanent resource such as a midden, mound or burrow. I conclude that there is no indication from Townsend's voles or any other microtines that females abandon their breeding range to their female offspring as a form of parental investment. Received: 27 February 1996 / Accepted after revision: 1 March 1997     

Reproductive activities in the sesarmid crab Sesarma intermedia in the coastal and estuarine habitats of Hakata,Japan

Aspects of activity pattern, mating, breeding behaviour and reproductive organs were studied in Sesarma intermedia (de Hann) along the Tatara river estuary, Hakata Bay, western Japan. Activity started from early May after a six-month hibernation period. The daily activity pattern was bimodal with one peak after sunset and another before sunrise. Mating started after emerging from hibernation and was ended by July. Practically no mating was seen in August and thereafter, by which time most of the females were carrying eggs. Larger males mated more frequently with any sized adult female. Copulation seemed to take place both in the burrow and on the surface of natural shelter. Females began to carry eggs from mid June on the estuarine habitat, whereas on the coastal habitat ovigerous females could be seen from mid-May. Multiple ovipositions of eggs were realized. Females were able to oviposit three times without any repeated mating. It took about 30 d to incubate one egg mass and less than one week to spawn again. The release of larvae was not sporadic but rather took place within a limited time interval. Sunset seemed to have a great influence on larvae release along the seashore. Energy allocation for reproductive purposes seemed to start before crabs awoke from the hibernation period. Towards the end of the breeding period there was a drastic decline in ovarian development with a remarkable gain in hepatopancreas, a reservoir for energy.     

Role of sex and breeding status in grooming and total tick load of impala

The lesser mouse lemur (Microcebus murinus) is a prosimian primate which presents evidence of sex ratio bias of offspring that agrees with the direction of bias predicted by the local resource competition model for facultative sex ratio adjustment. That is, females overproduced sons when grouped prior to mating, whereas isolated females exhibited the opposite tendency. In this solitary species, social communication relies heavily on urinary chemical signals. To test the hypothesis that sex biases induced by social factors may be linked to urinary cues, isolated females were exposed (n = 76) or not (control group, n = 16) to urinary cues from other reproductively active females from the beginning of the breeding season (induced by long photoperiod) until oestrus. During that period, females were either continuously (n = 17) or partially (n = 59) exposed to chemosignal stimulation. Females in oestrus were placed in contact with a breeding male and subsequently isolated until they gave birth. All females entered oestrus but the timing of oestrus was significantly delayed by 1 week in urine-exposed females. A general depressive effect of long-term urine exposure on fecundity was demonstrated, involving fewer impregnations, higher abortion frequency and smaller litter sizes. Among females giving birth (n = 55) to a total of 129 young, a significant positive correlation was found between sex ratio bias towards males and the duration of urine exposure. However, the shift in sex ratio at birth depended on the duration of urine stimulation during a sensitive period extending from the beginning of the long photoperiod until the beginning of the follicular phase. In the absence of urinary cues during the sensitive period, females significantly overproduced daughters (32% males of 53 newborn). As urine exposure increased during the sensitive phase, the proportion of males in litters increased from 54% males (n = 50) in partially urine-exposed females to a significant bias towards males (69.2% of 26 newborn) in totally exposed females. The biased sex ratio in response to chemical cues did not show consistent relationships with maternal body weight, parity or litter size. Although the intrinsic mechanisms involved in sex-biased conceptions are not known, chemical cues could interact with the photoperiodic control of gonadotropin secretions. Received: 14 January 1995/Accepted after revision: 26 November 1995