Inter-litter competition and communal nursing among Norway rats: advantages of birth synchrony

Summary The effect of inter-litter competition on pup survival was investigated in pairs of female rats (Rattus norvegicus) living and breeding in the same environment. If a female gave birth when a 0- to 14-day-old litter was already present in the environment, her pups had a very high chance of surviving, similar to the situation in which no other litter was present. Moreover, the mother was likely to nurse communally with the mother of the 0- to 14-day-old litter. This communal nursing benefitted the newborn pups as evidenced by their being heavier at weaning than litters that were not nursed communally. In contrast, if a female gave birth when a 15-to 28-day-old litter was already present in the environment, her newborn pups were likely to die within 3 days postpartum, owing to the fact that they were often prevented from suckling at their mother's teats, resulting in milk deprivation, and were often beneath the older pups, resulting in physical trauma. These findings suggest that inter-litter competition is an important source of pup mortality when litters are born 15-28 days apart. The data are discussed in terms of the advantages of birth synchrony.     

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.     

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.     

Sex ratio shifts within litters of meadow voles (Microtus pennsylvanicus)

Summary Changes in the sex ratio of juvenile recruits into a population of meadow voles (Microtus pennsylvanicus) were correlated with shifts in the weight and mortality of pups within the population. The biased recruitment of female juveniles in the spring was reflected in differential allocation of energy within the litters, as measured by female pups being heavier than male pups (n=245). In the fall, the shift in recruitment to male juveniles was reflected within litters by male pups being heavier than female pups (n=139). Nestling mortality showed a similar gender bias. Skewed sex ratios were most evident within the litters of larger mothers, indicating the gender bias was not trigered by energy limitations. We postulate that gender differences in social spacing and behavior result in spring/fall fluctuations in the reproductive success of offspring, based on their gender.     

Influence of social factors on sex ratio at birth,maternal investment and young survival in a prosimian primate

Summary In order to determine whether social factors influence sex ratio at birth in lesser mouse lemurs, experiments were conducted during 5 successive breeding periods on 51 females. At the beginning of the breeding season, females were either isolated (I) or grouped (G) in heterosexual groups with an increasing number of females (2, 3 or 4). To ensure mating, I females were introduced in a group only during the oestrous period. After mating, both I and G females were isolated during pregnancy and lactation. Reproductive capacities of females in terms of oestrus occurrences (n = 324), impregnations (n–210), pregnancies (n = 136) or abortions (n = 38) or litter sizes (1–3 young) were affected neither by age and parity of females nor by group housing prior to conception. G females produced significantly more sons than daughters (67% males for 189 newborn) while females living alone except during the mating period demonstrated a significant inverse tendency (39.6% males for 96 newborn). Distribution of sexes in litters was statistically different from random and varied according to the shift of sex ratio at birth. In G females, the shift in the sex ratio towards males was consistent across the different groups, independent of the number of females living together, suggesting that the presence of only 1 female is sufficient to induce a bias in the sex ratio. No correlation was found between infant survival at weaning and age, parity or group housing of the mother. The maternal investment allocated to male or female newborn was similar provided the litter contained at least 1 male. In litters without males, growth and survival of female infants were significantly less. These results on sex ratio bias in captive female mouse lemurs agree with directions of bias predicted by the local resource competition model for facultative sex ratio adjustment (Clark 1978). Nevertheless, the pattern observed in mouse lemurs appears to be independent of the nutritional state of the female and of differential maternal investment.     

Evidence for obligate monogamy in the Djungarian hamster,Phodopus campbelli: pup survival under different parenting conditions

Summary Two recent studies have described pregnancy blocking responses in female Djungarian hamsters, Phodopus campbelli, under conditions where the potential for paternal investment by the male is low (Wynne-Edwards and Lisk 1984; Wynne-Edwards et al. 1987). If this pattern of pregnancy block is adaptive, solitary females should have a low probability of successfully rearing a litter without investment by their mate. The present study was designed to compare the relative reproductive success of solitary females with that of paired females. Weights and survivorship were determined daily from birth through weaning (day 18 after birth) for litters of Djungarian hamster pups raised by solitary females (Solitary females), male-female pairs (Pairs), or female-female pairs consisting of the experimental female and her unmated littermate sister (Sisters). Unmated littermate sisters were included in the design to differentiate between the contribution of the mate and the contribution of another conspecific adult to the reproductive success of the experimental female. As predicted by the hypothesis that the pattern of incidence of pregnancy block was adaptive (Wynne-Edwards et al. 1987), the presence of the mate significantly affected the reproductive success of the female. Pairs were very successful, raising 95% of pups and 100% of litters to weaning. Solitary females were significantly less successful, raising only 47% of pups and 77% of litters with 16% of litters intact on day 18. A littermate sister did not compensate for the absence of the mate. Sisters raised only 61% of pups and 73% of litters with 36% of litters intact on day 18. Pup survival and weights were independent of sex. However, for pups that were still alive on day 18, weight distributions did not differ significantly across the three parenting conditions. Likewise, the proportion of best pups alive on day 18 (with weights exceeding a threshold of 11.5 g) did not differ significantly across groups. Differences between parenting conditions lay in the survivorship of pups rather than in the quality of the pups that survived. This study provides evidence that the presence of the mate is essential for high female reproductive success even under the ideal climate and nutritional conditions of the laboratory. Thus, the Djungarian hamster is a strong candidate for a small mammalian species with obligate monogamy as the optimal mating system.     

Evidence of equal maternal investment in the sexes in the polygynous and sexually dimorphic grey seal (Halichoerus grypus)

In polygynous and sexually dimorphic mammals, parents may be expected to bias their investment towards sons because variation in reproductive success is usually higher among males than among females. Moreover, male reproductive success often depends on adult body size, which, in turn, may depend on the level of parental investment. We therefore predicted that in the grey seal (Halichoerus grypus), a polygynous and sexually dimorphic phocid seal, females should invest more in individual sons than in individual daughters. We found that male pups were born heavier than female pups, but that the growth rates and suckling behaviour were similar for the two sexes. The growth rates and the birth weights were not correlated for the pups of either sex. Mothers did not behave differentially towards offspring of the two sexes, except that mothers of male pups spent more time in visual contact with their pups. Male and female pups had similar activity levels and begged at similar rates. We argue that reports of equal expenditure on the two sexes can be accepted as evidence of equal investment, provided that three assumptions are fulfilled. First, parental care must be costly to the parent. Second, energy expenditure must be the most important component of parental investment. Third, there must be no negative correlation between maternal body condition and the ratio of sons to daughters produced. We argue that these assumptions are met in our study, and that our results provide evidence of equal maternal investment in the sexes in grey seals.     

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     

Parental investment in southern elephant seals,Mirounga leonina

Summary The southern elephant seal is among the most sexually dimorphic and polygynous of all mammals: males may be more than 10 times the weight of reproducing females and only the largest 2–3% of males are likely to breed. Current optimization theories of sexual selection predict that evolution would favor greater parental investment in individual males than in females. Because southern elephant seals represent an extreme of polygyny and sexual dimorphism, they might be expected to show a dramatic difference in parental investment in male and female pups. However, in a study of parental investment in elephant seals at South Georgia, using several different methods, we found no such difference after parturition. Mother-pup pairs were immobilized and weighed early in lactation, recaptured near the end of lactation and reweighed. A further 30 pups were weighed an average of five times during lactation to establish the shape of the growth curve and to serve as partial controls for the previous set of animals. Initial post-partum weight in females ranged from 346 to 803 kg (=506, SD=111, n=26). Pup birth weight was related to mothers' post-partum weight in female pups but small females often gave birth to large male pups. Male pups were significantly heavier at birth than females. However, this size difference did not persist. Male and female pups were suckled for the same period, grew at the same rate and were not significantly different in weight at weaning. Mothers lost weight at the same rate regardless of their pup's sex.     

Sperm competition in deer mice (Peromyscus maniculatus bairdi)

Summary Six experiments were conducted to determine the effects of mode of estrus (cycling versus postpartum) and intervals between matings on sperm competition in deer mice. When there was no lengthy interval between matings by different males, sperm competition matings by different males, sperm competition with females in postpartum estrus was the same as that with females in cyclin estrus (i.e., no male order effect). Imposition of 4 h delays between males did not engender a mating advantage when females were in postpartum estrus. When 4 h intervals were imposed with females in cycling estrus and tests were begun relatively early in the dark phase, there was a secondmale advantage with respect to the number of pups sired. When minimal delays were imposed with females in cycling estrus the time of testing within the dark phase did not affect the pattern of sperm competition, but larger litters were born following matings later in the dark phase. When 4 h delays were imposed with the first male mating late in the dark phase a first-male advantage resulted. It is hypothesized that for females in cycling estrus there is a period of peak fertilizability. There is no straight-forward effect of delays between males per se; rather, effects must be analyzed in relation to the period of peak fertilizability. The timing of insemination appears less critical for postpartum females.     

Aggression and submission reflect reproductive conflict between females in cooperatively breeding meerkats Suricata suricatta

In many cooperatively breeding species, dominant females suppress reproduction in subordinates. Although it is commonly assumed that aggression from dominant females plays a role in reproductive suppression, little is known about the distribution of aggressive interactions. Here, we investigate the distribution of aggressive and submissive interactions among female meerkats (Suricata suricatta). In this species, dominant females produce more than 80% of the litters, but older subordinates occasionally breed. Dominant females commonly kill the pups of subordinates and usually evict older female subordinates from the group 1–3 weeks before the birth of the dominant female's litter. The aggression frequency of the dominant female toward subordinates and the submission frequency that each subordinate female showed to the dominant female increased as the age of the subordinate female increased and as the birth of the dominant female's pups approached. Moreover, as birth approached, both of these behaviors intensified more quickly between the dominant female and older subordinates than between the dominant female and younger subordinates. The aggression frequency of the dominant female toward each subordinate female predicted whether that subordinate female was evicted from the group; the submission frequency by each subordinate female predicted the timing of their eviction during the pregnancy period of the dominant female. These results support the idea that conflict between dominant and subordinate females increases with the age of subordinates and, since older subordinate females are most likely to reproduce, suggest that dominant females may less easily control reproductive attempts by older subordinate females.     

Does male harassment of females contribute to reproductive synchrony in the grey seal by affecting maternal performance?

We investigated the possibility that male harassment of lactating females differed in relation to time of birth in the grey seal, Halichoerus grypus, on Sable Island, Nova Scotia. This was done by comparing the frequency of male disturbances, maternal performance and pup growth for females that either gave birth during the peak of the pupping season or after the peak. Of the females, 58% gave birth in a 7-day period near the beginning of the pupping period, when the operational sex ratio was 2–4 females per male. Late in the pupping period the operational sex ratio reversed to about 1 female for every 2 males. The relative frequency of disturbances by males was significantly greater for late-pupping mothers than for peak-pupping ones (1.9 vs. 1.4 encounters/h). Females that gave birth late also were disturbed by males 3 times more often than females that gave birth during the peak (3.4 vs. 1.1 % of observation time). Late-pupping mothers spent 22% less time suckling (4.0 vs. 5.1 % of observation time), had 30% slower growing pups (1.7 vs. 2.4 kg/d), and weaned pups that were 16% lighter (45.6 vs. 54.0 kg). The effect of birth time on pup mass gain and weaning mass was not attributable to factors such as maternal mass, pup birth mass or pup sex. We conclude that the reduced maternal performance is likely the result of the increased male harassment. As reduced weaning mass can lead to reduced juvenile survival, male harassment may have contributed to the enhanced reproductive synchrony in this species.     

Offspring sex ratio in relation to female size in southern elephant seals,Mirounga leonina

Southern elephant seals Mirounga leonina display extreme sexual dimorphism. In addition females show great variation in size and stored resources at parturition. Therefore they present an excellent opportunity for examination of responses of sex ratio to resource availability. We studied the relationships between the size of southern elephant seal females at parturition and the size and sex of their pups at South Georgia over four breeding seasons. We found a large individual variation in maternal post-partum mass (range 296–977 kg, n=151). Larger mothers gave birth to larger pups, irrespective of the sex of their pup. Male pups were on average 14% larger than females at birth and consequently more costly to bring to parturition. Our results suggest that female southern elephant seals must weigh more than 300 kg if they are to breed at all, and more than 380 kg if they are to give birth to a male pup. Above this threshold the proportion of males among offspring rapidly increases with maternal mass, and stabilizes at a level not significantly different from parity. These results show that smaller females of southern elephant seals vary offspring sex ratio in a way that is consistent with theories on adaptive offspring sex ratio. A smaller mother with a male foetus may benefit from terminating her pregnancy and allocating the resources she saves to her own growth. She could then give birth to and raise a larger pup in the subsequent season.     

Sexual dimorphism in sea lion pups: differential maternal investment,or sex-specific differences in energy allocation?

Proximal mechanisms underlying a faster growth rate in male compared to female California sea lion pups were investigated. Males are significantly larger at birth than females. Specifically, we asked if differential maternal investment contributed to enhanced male growth via: (1) larger mothers having disproportionately more male pups, (2) more time and energy put into foraging by mothers of male pups, and (3) greater milk production in mothers of male pups. We also considered four aspects of differential energy utilization and acquisition by male and female pups: (1) male pups attempting to save energy for growth by changes in behavior, (2) longer suckling bouts with mother and more sneak suckling of non-filial females by male pups, (3) lower maintenance costs in males via a lowered resting metabolic rate, and (4) increased assimilation efficiency in males. Our study showed that there are no differences in the size of females or length of foraging trips for mothers of male and female pups. Male pups received more milk from their mothers, but the difference was no longer significant when the larger body size of males was considered. There were no differences in either the activity budgets or suckling behavior of male and female pups. Male pups, however, did have lower resting metabolic rates than females. We conclude that enhanced male perinatal growth is a consequence of a larger size at birth, proportionally more milk from mothers to support the greater demands of larger body size, and lower maintenance costs due to a lower resting metabolic rate. Received: 28 April 1995/Accepted after revision: 25 July 1995     

Sex ratio and maternal rank in wild spider monkeys: when daughters disperse

Summary Data from a long-term field study of the spider monkey, Ateles paniscus, in Peru indicate that a strongly female-biased sex ratio exists from birth in this population. Of 46 infants born between July 1981 and June 1986, 12 were male, 32 were female and 2 were of undetermined sex. This effect is consistent between years as well, with more females than males born in each year of the study (Table 1). This bias is driven by the fact that low-ranking females produce daughters almost exclusively, while high-ranking females bias their investment somewhat less strongly towards sons (Table 2). The unusual pattern of female-biased maternal investment observed in this population of Ateles probably occurs for a combination of the following reasons: (1) maternal investment in individual male offspring is somewhat greater than in individual female offspring; (2) males remain with their natal groups, and the sons of high-ranking females are likely to be competitively superior to the sons of low-ranking females; (3) males compete for mates, and only the one or two most dominant males within a community are likely to achieve significant reproductive success. Two possible mechanisms of sex-ratio adjustment and the evidence for each are discussed.     

Offspring sex ratios reflect lack of repayment by auxiliary males in a cooperatively breeding passerine

The repayment hypothesis posits that primary sex ratios in cooperative species should be biased towards the helping sex because these offspring “repay” a portion of their cost through helping behavior and therefore are less expensive to produce. However, many cooperatively breeding birds and mammals do not show the predicted bias in the primary sex ratio. Recent theoretical work has suggested that the repayment hypothesis should only hold when females gain a large fitness advantage from the presence of auxiliary adults in the group. When auxiliaries provide little or no fitness advantage, competition between relatives should lead to sex ratios biased towards the dispersing (non-helping) sex. We examined the benefits auxiliaries provide to females and corresponding offspring sex ratios in the red-backed fairy-wren (Malurus melanocephalus), a cooperatively breeding Australian bird with male auxiliary helpers. We found that auxiliaries provide little or no benefit to female reproductive success or survival. As predicted, the population primary sex ratio was biased towards daughters, the dispersing sex, and females with auxiliaries produced female-biased broods whereas females without auxiliaries produced unbiased broods. Moreover, offspring sex ratios were more strongly biased toward females in years when auxiliaries were more common in the population. These results suggest that offspring sex ratios are associated with competition among the non-dispersing sex in this species, and also that females may use cues to assess local breeding opportunities for their offspring.     

Reproductive success in a low skew, communal breeding mammal: the banded mongoose, Mungos mungo

In most cooperatively breeding species, reproduction is monopolised by a subset of group members. However, in some species most or all individuals breed. The factors that affect reproductive success in such species are vital to understanding why multiple females breed. A key issue is whether or not the presence of other breeders is costly to an individual’s reproductive success. This study examines the factors that affect the post-parturition component of reproductive success in groups of communal-breeding banded mongoose (Mungos mungo), where up to ten females breed together. Per-litter reproductive success was low (only 18% of pups survived from birth to independence). Whilst singular breeding was wholly unsuccessful, there were costs associated with breeding in the presence of increasing numbers of other females and in large groups. Synchronisation of parturition increased litter success, probably because it minimises the opportunity for infanticide or decreases competitive asymmetry between pups born to different females. There was no evidence of inbreeding depression, and reproductive success was generally higher in litters where females only had access to related males within their group. I conclude that communal breeding in female banded mongooses represents a compromise between the benefits of group-living and communal pup care on the one hand, and competition between females to maximise their personal reproductive success on the other. Such conflicts are likely to occur in most communal breeding species. Whilst communal breeding systems are generally considered egalitarian, negative effects of co-breeders on individual reproductive success is still an issue.     

Allocation of reproductive effort in Mus domesticus: responses of offspring sex ratio and quality to social density and food availability

Summary Female mammals in good condition can maximize their inclusive fitness by investing more in male offspring than in female offspring during periods of poor environmental quality. To test this hypothesis, we measured the effects of undernutrition and crowding before and during gestation on the sex ratio and weight of offspring at parturition and at weaning in Mus domesticus. Sex ratio was not significantly affected by density. Dams altered the sex ratio of their offspring in response to food availability, but only if variance in competitive success within the experimental subpopulation was evident. Thus ad lib fed females produced litters with an unbiased sex ratio, competitively successful females under moderate food availability produced a male-biased sex ratio, and severely food deprived females produced litters with a female-biased sex ratio. In groups that experienced competition for food, successful dams favoured male offspring during lactation. These results are consistent with the predictions of Trivers and Willard (1973). Analysis of within-cell variance and covariance suggests that the interaction of social structure and food availability provides specific cues for the dams' tactical reproductive choices.     

Infant care,infanticide, and female reproductive strategies in polygynous groups of common marmosets (Callithrix jacchus)

Reproduction in marmoset and tamarin groups is typically restricted to a single dominant female, but it is unclear why subordinate females tolerate delayed reproduction. The presence of two breeding females in free-ranging groups of common marmosets (Callithrix jacchus) presented a unique opportunity to examine differences in the reproductive strategies of dominant and subordinate females. Three groups were monitored for 12–18 months at a forest reserve in northeastern Brazil. Data on infant care were collected during two consecutive all-day follows every 10 days until the infants were 2 months old. Carrying patterns for infants born to dominant females were similar to those observed in groups containing a single breeding female. All group members over 5 months of age participated in infant care, and dominant females allowed some group members to carry their infants from the 1 st day of life. In contrast, subordinate females were protective of their offspring and were their sole caretakers for at least a week following birth. One infant born to a subordinate was killed in an attack involving the dominant female. Overall, dominant females gave birth to more infants and had higher infant survival than did subordinate breeding females. subordinate females were successful in rearing young only when the timing of births was such that they did not overlap with the dependency period of infants born to the dominant female. These patterns suggest that subordinate females may face resource competition, especially over access to helpers. In times of severe competition, subordinate females that delay reproduction may be avoiding a wasted reproductive effort.     

Sibling competition in guinea pigs (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): scrambling for mother’s teats is stressful

Lactation is the most energy-intense period in the life of a female mammal. This can cause severe conflict between mother and offspring over the duration of lactation but also between siblings over the amount of milk each pup gets from its mother. Thus, competitive interactions between siblings are expected, and competition is likely to increase with litter size, particularly in species where the number of offspring exceeds the number of teats. We studied sibling competition in the domestic guinea pig (Cavia aperea f. porcellus), which has two teats, but frequently bears litters of up to five pups. By cross-fostering we created non-competition (control) litters with two pups and competition litters with four pups and observed nursing behaviour on days 5, 10, 15 and 20 postpartum. Pups of larger litters had lower growth rates, indicating increased competition among siblings in these litters. Pups of larger litters had to wait longer for access to a teat and spent less time suckling than pups of smaller litters but ate more solid food instead. Additionally, we manipulated the individual short-term need of pups by separating half of the pups of each litter for 2 h from their mothers before observation. Within a litter, hungry pups achieved access to milk faster and spent more time suckling than non-hungry pups. Pups competed mostly by scramble competition. Aggressive interactions occurred only in large litters. Pups of large litters had higher cortisol levels than pups in small litters. These effects decreased with age as pups became increasingly independent of maternal milk. Pup behaviour appears to fit better with models of scramble competition than with those of honest signalling. This contribution is part of the special issue “Sibling competition and cooperation in mammals” (guest editors: Robyn Hudson and Fritz Trillmich).