Age,coloration and dominance in nonbreeding hummingbirds: A test of the asymmetry hypothesis

Summary During the nonbreeding season, adult Anna and black-chinned hummingbirds (Calypte anna and Archilochus alexandri) have lower defense costs and more exclusive territories than juveniles. Adult C. anna are victorious over juveniles in aggressive encounters, and tend to monopolize the most temporally predictable resources.Juveniles are more successful than adults at stealing food from territories (the primary alternative to territoriality), presumably because juveniles are less brightly colored. Juveniles have lighter wing disc loading than adults, and consequently should have lower rates of energy expenditure during flight. Reduced flight expenditures may be more important for juveniles because their foraging strategy requires large amounts of flight time. These results support the contention of the asymmetry hypothesis that dominance can result from a contested resource being more valuable to one contestant than to the other.Among juveniles, defence costs are also negatively correlated with age and coloration; amount of conspicucus coloration is negatively correlated with the number of bill striations, an inverse measure of age.     

Dominance in feeding territories relates to foraging success and offspring growth in brown skuas <Emphasis Type="Italic">Catharacta antarctica lonnbergi</Emphasis>

Territoriality should lead to strict dominance, as territory holders typically control access to resources and exclude others from their use. In feeding territories, dominance should be reflected in foraging success and ultimately in reproduction differences; however, these successive links have rarely been made explicit. Therefore, we investigated a population of brown skuas Catharacta antarctica lonnbergi, in which only part of the breeding population occupied feeding territories within penguin colonies. We identified the dominance hierarchy and determined the foraging success of the participants in fights for access to penguin carcasses within the territories. Furthermore, we monitored offspring growth from parents with and without feeding territories. Our results indicated a clear dominance hierarchy with territorial birds in their own territory dominating over territorial breeders from other territories, non-territorial breeders and non-breeding birds. However, territory owners could not completely exclude others from access to food. Foraging success was positively related to dominance scores: The dominant territory owners received 63% of a carcass, whereas non-territorial pairs could get less than 10%. The link between foraging success and offspring development was less clear: Although male chicks of non-territorial parents suffered from lower growth rates and, thus, delayed fledging, there were no such differences in female chicks. Territoriality in skuas did not imply a complete occupation of food, but guaranteed optimal growth conditions for offspring. Non-territorial individuals were forced to search for alternative resources, and the restricted access to the preferred food resulted in inferior conditions for offspring development, making this foraging strategy less rewarding.     

The influence of kinship on foraging competition in Siberian jays

Foraging competition in Siberian jay groups was examined in relation to dominance and kinship to determine whether juvenile offspring, by associating with adults, gained in food acquisition relative to juvenile immigrants. Members of the adult pair were dominant over juvenile cohort members and males were dominant to females, although an inter-sexual hierarchy, with male juveniles occasionally overlapping adult females, was suggested. Few competitive asymmetries were found between adults and retained offspring or adults and immigrant juveniles when they were competing for food together, but in kin and non-kin foraging groups, respectively. Male offspring visited the bait site more frequently than adult males, and female immigrants spent less time at the bait site than adult females. Under these circumstances, hoarding activities may limit the ability of alpha members to control resources. In mixed groups containing both juvenile offspring and juvenile immigrants, no difference was found in the number of visits made to the bait site, although load sizes and foraging rates were lower for immigrant birds. Retained juveniles obtained greater load sizes and foraging rates when associating with adults. The social dominance of parents suggests that they control juvenile foraging. Although offspring benefit in the presence of adults, adults may incur a cost to their restraint by spending more time at the bait site when competing with immigrants. These results extend conclusions from previous work describing the role of selective tolerance by adults which relaxes competition with retained offspring in Siberian jay winter groups. The present findings suggest that offspring benefit in both immediate and future energy gains, which may have a direct influence on survival. Received: 18 September 1996 / Accepted after revision: 26 January 1997     

Trace metals in mussels (Mytilus edulis) from southwest Iceland

The temperate population of the tropical anemonefish Amphiprion clarkii has a seasonality of reproduction, larval settlement and growth. There was a considerable difference in size (15–47 mm) among 0-year olds in December, after the season of larval settlement and growth. This difference could be attributed mainly to a difference in the duration of the first growing season spent after settlement and a social inhibition of growth of the subordinate late settlers by frequent attacks by the dominant early settlers. The growth equation which was used to describe the growth of tagged individuals estimated that an anemonefish would get an opportunity to spawn within four of five years after settlement. Breeding adult pairs occupied almost all sea anemones essential to breeding within a 50×50 m study area, and all 17 new adults that matured from non-breeding juveniles to breeding adults between June 1983 and August 1985 were found in the territories from which either or both of the mated adults had disappeared. The new adults were large juveniles who had resided near those vacant territories before. These results suggest that the onset of breeding by an individual is not only determined by his age, but also by his ranking in the dominance hierarchy. Therefore, the difference in size among 0-year olds in December might give rise to the difference in age of the earliest breeding and might be a factor in making a female apply more reproductive effort at the beginning of the season.     

The effect of reproductive condition on the foraging behavior of female hoary bats,Lasiurus cinereus

Summary Female mammals experience larg changes in time and energy budgets associated with reproduction and these may influence the foraging strategies of individuals. I studied the changes in foraging behavior associated with reproduction in female hoary bats, Lasiurus cinereus. As lactation progressed, individuals departed to forage earlier in the evening and spent more time foraging per night and less time roosting with their young. Foraging time increased by at least 73% between early lactation and fledging and then declined as the young became independent. Females with two young foraged for longer than did those with one and females with pre- and postfledging young foraged in different habitats. The changes in foraging time suggest that foraging activity of female L. cinereus is constrained and individuals act as time minimizers, adjusting their foraging behavior to meet current energy demand. Predation risk is unlikely to constrain the behavior of these bats. However, maximizing energy intake throughout lactation may not be the optimal strategy because storing excess energy increases flight cost and may reduce foraging efficiency. The need to keep newborn young warm may also influence foraging time. Such constraints, causing changes in foraging activity, may alter the availability of habitats and prey and must be considered when modelling foraging strategies. In addition, changes in flight time may significantly alter the energy budgets of bats in different stages of reproduction.     

Social behavior of free-ranging juvenile sooty mangabeys (Cercocebus torquatus atys)

Sooty mangabeys are terrestrial monkeys exhibiting female philopatry and male dispersal. Studies in captivity as well as in the wild have found that adult females form linear dominance hierarchies. However, while captive studies found no evidence for a matrilineal social system, a previous study in Taï National Park, Ivory Coast, suggested that relatedness could influence both dominance rank and affiliation pattern among adult females. Here I test whether the dominance rank, coalitionary behavior, and affiliative behavior of juveniles in a group of free-ranging mangabeys in the Taï National Park are in accordance with a matrilineal, individual, or age-related dominance system. I found that juvenile females' dominance ranks remained stable over time and were highly correlated with the dominance ranks of their mothers, whereas juvenile males' dominance ranks were initially correlated with the ranks of their mothers, but showed greater instability with increasing age. Moreover, coalitions occurred mainly between juveniles and animals that were close in rank, including their mothers and siblings. Finally, juvenile females associated and groomed preferentially with close-ranking juvenile and adult females. Juvenile males showed similar preferences in affiliation with adult females, but when associating with juvenile males, they preferred peers. The observed social behavior of free-ranging juvenile sooty mangabeys resembled the social behavior described for juveniles of many matrilineal primate species.     

Family structure and variation in reproductive success in blackbirds

In avian families, some offspring are rendered unequal by parental fiat. By imposing phenotypic handicaps (e.g., via asynchronous hatching) upon certain of their offspring and not others, parents structure the sibship into castes of advantaged “core” offspring and disadvantaged “marginal” offspring that results in an asymmetric sibling rivalry. Here, I show how this family structure scales up to population level reproductive consequences. In a 17-year study of red-winged blackbirds (Agelaius phoeniceus), I show that year-to-year variation in the number of surviving offspring is driven primarily by variation in the number of marginal offspring at hatching and their posthatching survival. Clutch size, core brood at hatching, and fledging varied little from year to year and had little direct effect on year-to-year variation in total brood size at fledging; conversely, variation in the size of the marginal brood at hatching and at fledging was much greater. Marginal but not core brood size at hatching rose with mean clutch size; in years where parents laid larger average clutches they did so by adding marginal progeny. The mean posthatching survival of marginal offspring was always lower than that of core offspring in a given year, and there was no overlap in the distributions. The highest mean survival of marginal offspring across years fell below the lowest mean survival of core offspring; broods were deeply structured. There was an overall female bias among fledglings, and the sex ratio varied across years, with a higher proportion of the smaller female nestlings in years of below average reproductive success. Such variation was especially pronounced in the marginal brood where a higher incidence of brood reduction allowed greater potential for sex-biased nestling mortality. In years of the highest average reproductive success, the sex ratio in the marginal brood approached equality, whereas in years of the lowest average reproductive success, more than two thirds of 8-day-old nestlings were female. Structuring the brood into core and marginal elements allowed parents to modulate both offspring number and sex under ecological uncertainty with direct consequences for population-level reproductive success. They produced fewer and less expensive fledglings in below average years and more and more expensive fledglings in above average years.     

Kinship and social tolerance among female and juvenile wild house mice: kin bias but not kin discrimination

Kin-biased social tolerance among house mice has been interpreted in terms of kin discrimination. However, several lines of evidence suggest it may instead be an incidental artifact of group member discrimination. This leads to very different predictions about the social consequences of relatedness within and between social groups. Social interactions between wild-stock adult female and juvenile house mice (Mus domesticus) established in neighbouring territorial groups within enclosures reveal relatedness to dominant males within groups as the major factor determining social tolerance of juveniles by females. Relatedness to the female herself had no significant independent effect on responses indicating tolerance. Females were generally more aggressive toward neighbouring-group juveniles (all unrelated to females) compared with those from their own group (all related to females), but were most aggressive toward neighbouring juveniles sired by the neighbouring dominant male. They were also more aggressive toward their own-group juveniles that had been sired by the neighbouring dominant but only when encountered in the neighbouring territory and with a greater bias against female juveniles. Females were least aggressive toward own-group juveniles sired by their own-group dominant male. The sire-bias in tolerance among females is similar to that reported among the dominant males themselves in an earlier study. As a result of the combined sire-bias in tolerance by adult males and females, juveniles sired by their own-group dominant males become less likely to intrude into a neighbouring territory with time. Overall, the results suggest that differences in social tolerance reflect discrimination on the basis of social group membership rather than relatedness between interactants and thus provide strong experimental evidence in support of incidental kin bias rather than kin discrimination.     

The superseded female's dilemma: ultimate and proximate factors that influence guarding behaviour of the carpenter bee Xylocopa pubescens

Both solitary and primitively social nests of the facultatively social carpenter bee Xylocopa pubescens can be found throughout most of the breeding season. In social nests there is reproductive division of labour between a dominant forager and a guarding female. Two types of guarding females can be discerned: the young pre-reproductive guards, and older, formerly reproductive guards. The latter type of guard is found when, after a take-over of reproductive dominance either by a nestmate (mostly a daughter) or an intruder, the defeated female stays in the nest instead of leaving to try and found or usurp another nest. She is then manipulated into the role of a guard. The dominant female profits from the presence of the guard since she protects the nest against pollen robbery by conspecifics (Hogendoorn and Velthuis 1993). We have studied why superseded females might prefer to remain as a guard, rather than try their luck somewhere else. The hypotheses investigated pertain to (1) the difficulty for the defeated female of finding a new nest and of restarting reproductive activities due to (a) ecological constraints (nest and pollen shortage) and (b) the effect of age and wear on the defeated female; (2) the effects of guarding in terms of inclusive fitness. We found that superseded females remained as guards significantly more often when a nestmate (not necessarily close kin) took over reproductive dominance than when an intruder did so. Other factors associated with the decision of the defeated female to stay or leave were her age and the number of her own young still present after the supersedure. The probability of finding or constructing a new nest was lower for old than for young females. After finding a nest, old females produced less brood than young foundresses. As a result of these two factors old superseded females gained, in terms of inclusive fitness, by staying as guards, whereas young females profited from leaving the nest. We interpret these results as an indication that guarding behaviour has evolved due to kin selection. However, kin discrimination apparently did not occur. Therefore we conclude that in this species kin selection is not, in the proximate frame of reference, based on kin recognition and preference for helping kin. Correspondence to: K. Hogendoorn     

Fitness consequences of within-brood dominance in the cooperatively breeding acorn woodpecker

Social dominance confers potential advantages in terms of access to superior resources, habitats, and breeding opportunities. In the cooperatively breeding acorn woodpecker (Melanerpes formicivorus), within-brood dominance among juveniles is correlated with relative body size as nestlings. Capitalizing on this relationship, we investigated the fitness consequences of dominance by means of paired comparisons of broodmates. We found that (1) larger fledglings retained at least some of their size advantage as adults; (2) overwinter survival of larger, dominant fledglings was significantly greater than subordinates, but was not relatively greater when resources were poor than when they were good; (3) among birds surviving their first winter, there were no differences vis-à-vis dominance in terms of the proportion of birds acting as helpers or inheriting their natal territory. However, larger, dominant males were present in the study area longer than subordinates, suggesting that they either survived better or were more successful at gaining reproductive opportunities; (4) if only one male broodmate became a helper instead of dispersing, he was significantly more likely to be the smaller subordinate, consistent with the view that helping is a best-of-a-bad-job strategy; and (5) there were no significant differences in reproductive success among pairs of male broodmates that cobred together as adults, consistent with prior work failing to detect a phenotypic correlation of reproductive skew. Our results indicate that within-brood dominance relationships established as juveniles have significant effects on first-year survivorship and at least some aspects of adult fitness.     

Dominance behaviour,body weight and fat variations,and partial migration in European blackbirds Turdus merula

Summary A partially migratory population of European blackbirds Turdus merula was studied in central Europe during the non-breeding season. Body weight and the amount of subcutaneous fat deposits showed seasonal variations typical for each age and sex category. Juveniles were fattest in autumn and spring and both juveniles and females lost fat and body weight in mid-winter. Adults and males reached their maximum body weight in mid-winter. These findings together with data from Schwabl (1983) suggest that juveniles and females constitute the main fraction of those birds migrating from the area in autumn. In winter mainly adults and males have access to food resources. These patterns may be regulated by dominance behaviour, since adults and males are usually dominant over juveniles and females throughout the non-breeding season. Subordinate birds that stay in the area probably suffer from a higher winter mortality but have the advantage of early breeding.     

Dominance,prior occupancy,and winter residency in the great tit (Parus major)

Summary This study reports an aviary experiment aimed at determining what affects social dominance in the great tit (Parus major), especially why older birds (adults) in nature normally dominate younger ones (juveniles). When birds were matched with respect to age, prior residency determined dominance. Without a difference in prior residency older birds dominated younger ones. However, when juvenile birds had a prior residency advantage over adult birds, they often became dominant. This was especially so when the juvenile bird was large relative to the adult bird. When a resident juvenile male was also consorted by a female, he became dominant over an adult male on most occasions. An experiment where the dominant bird was removed and later returned to the aviary failed to produce more than one shift in dominance. However, the proportion of reversals in dominance interactions increased with separation time. It is argued that the fact that dominance depends on prior residency selects for winter residency in the great tit.Offprint requests to: H.G. Smith     

Maternal characteristics and the production and recruitment of sons in the eastern kingbird (<Emphasis Type="BoldItalic">Tyrannus tyrannus</Emphasis>)

Sex allocation theory predicts that if variance in reproductive success differs between the sexes, females who are able to produce high-quality young should bias offspring sex ratio towards the sex with the higher potential reproductive success. We tested the hypothesis that high-quality (i.e., heavy) female eastern kingbirds (Tyrannus tyrannus) that bred early in the breeding season would produce male-biased clutches. A significant opportunity for sexual selection also exists in this socially monogamous but cryptically polygamous species, and we predicted that successful extra-pair (EP) sires would be associated with an excess of male offspring. Although population brood sex ratio did not differ from parity, it increased significantly with female body mass and declined with female breeding date, but was independent of the morphology and display (song) behavior (correlates of reproductive success) of social males and EP sires. Male offspring were significantly heavier than female offspring at fledging. Moreover, the probability that male offspring were resighted in subsequent years declined with breeding date, and was greater in replacement clutches, but lower when clutch size was large. Probability of resighting female offspring varied annually, but was independent of all other variables. Given that variance in reproductive success of male kingbirds is much greater than that of females, and that male offspring are more expensive to produce and have a higher probability of recruitment if fledged early in the season, our results support predictions of sex allocation theory: high-quality (heavy) females breeding when conditions were optimal for male recruitment produced an excess of sons.     

Sexual selection in the northern pintail (Anas acuta): the importance of female choice versus male-male competition in the evolution of sexually-selected traits

We experimentally studied the relative importance of plumage, dominance status, and courtship behavior in determining male pairing success in the northern pintail Anas acuta and assessed whether these traits function in female choice, male-male competition or both. In an experiment (experiment IA) that eliminated the confounding effects of male-male competition and social courtship, females chose males with pure white breasts and colorful scapular feathers. When the same group of birds were free to interact (experiment 1B), male behavior was more important: females chose males that courted them intensely and were attentive to them, although preferred males again had whiter breasts and more colorful scapulars. In a second experiment (experiment 2), testing the effect of age on pairing success, females showed a significant preference for 2-year-old males over yearlings: 2-year-old males courted more and were more attentive to the female than yearlings; they were also more colorful than yearlings in a number of plumage measurements. Although males (in both experiments 1B and 2) were aggressive to one another while courting the female and dominant males were sometimes able to exclude subordinates from social courtship, contrary to expectation, we found no relationship between initial dominance rank and pairing success or dominance rank and age. In addition, dominance was not correlated with any of the morphological traits measured. Once chosen, however, subordinate males typically initiated fights with the higher-ranked male(s) and quickly achieved dominance. These results suggest that (1) females choose males based on a suite of morphological and behavioral characteristics, (2) male dominance relationships do not constrain active female choice, (3) a male's position in a dominance hierarchy is largely a result rather than a cause of female choice, and (4) female choice plays a more significant role than male-male competition in the evolution of several secondary sexual traits in male northern pintails.     

Higher rate of nest loss among primary than secondary females: infanticide in the great reed warbler?

An individually marked population of great reed warblers Acrocephalus arundinaceus was studied between 1984 and 1991 in South Central Sweden. The fates of 279 nests were analysed for the 25 days following clutch initiation. The average frequency of nest loss (43%) did not differ significantly between years or between four periods of the breeding season. After a breeding failure, 70% of the females laid a replacement clutch, most of them together with the same male. On polygnous territories, males assisted the female who first hatched young regardless of her initial mating status (Fig. 2). According to the acutal status of the females attending the nests, for each day nests were classified as being of either monogamous (M), primary (P) or secondary (S) status. From egg-laying to fledging the rate of nest loss decreased among nests of primary status whereas it increased among nests of monogamous and secondary status. During the egg-laying period, the rate of nest loss was 3 times higher among nests of primary than among nests of monogamous and secondary status (Fig. 3). Thus, the high loss level among nests of primary status during the laying period was closely associated with the presence of a female with a less advanced nest on the territory. All nests were situated in reed beds above deep water and most of them at a height at which possible inter-specific nest predators would have caused disturbance to the nest itself. The suspicion that secondary females committed sexually selected infanticide was supported by an experiment with dummy eggs that revealed bill peck markings identical to those obtained from great reed warblers (Fig. 4).     

Sexual selection and mating system in Zorotypus gurneyi Choe (Insecta : Zoraptera)

Summary Social behavior of a species in the little-known insect order Zoraptera is described for the first time. Zorotypus gurneyi Choe (Insecta: Zoraptera) is a wing-dimorphic species that lives colonially under the bark of rotting logs in central Panama. Males are larger than females in total body size and fight each other to gain access to females. Highly linear and stable dominance hierarchies exist among males. Higher-ranking males show such agonistic behavior as jerking, chasing, head-butting, hindleg-kicking, and grappling, whereas subordinates often try to avoid contacts. Higher-ranking males, the dominant males in particular, are well recognized by others and relatively free of injuries. Although the dominant males are often the largest, the correlation between body size and dominance rank is not always significant. The mating system of Z. gurneyi is an example of female defense polygyny in which the dominant males obtain the majority of matings (75% on average). Mating success among Z. gurneyi males is much more variable than that of some lekking species.     

Temporary alteration of local social structure in a threatened population of Cuban iguanas (Cyclura nubila)

In small, insular populations, behavioral patterns that lead to increased variance in individual reproductive success can accelerate loss of genetic variation. Over a 1-year period, we documented behavior and hormone levels in a breeding group of adult Cuban iguanas (Cyclura nubila) at Guantánamo Bay. Male dominance was associated with body and head size, display behavior, testosterone levels, home-range size, and proximity to females. Based on their success in agonistic encounters, we ranked males in a linear dominance hierarchy. During the subsequent breeding season, we conducted a removal experiment in which the five highest-ranking males were temporarily relocated from the study site. Although we were unable to assess reproductive success directly, previously lower-ranking males assumed control of vacated territories, won more fights, and increased their proximity to females in the absence of the dominant males. When it results in greater mating opportunities for otherwise socially suppressed individuals, temporary alteration of local social structure may help limit erosion of genetic variation in small, insular populations. Electronic Publication     

Extra-pair paternity uncommon in the cooperatively breeding bicolored wren

We investigated parentage using multilocus DNA fingerprinting for 222 juveniles produced during 99 group-years in the bicolored wren Campylorhynchus griseus, a cooperatively breeding bird of the Venezuelan savanna. Young adult bicolored wrens (auxiliaries) remain in their natal territories and substantially enhance the production of young there. We have previously used behavioral indicators of dominance by a single male/female pair (principals) to infer breeding status, resulting in the commonly applied model of helping in which current fitness accrues to auxiliaries only indirectly, in proportion to their relatedness to the principals and the effect of their assistance on breeding success. Our parentage analysis has demonstrated that 8.6% of the juveniles found on territories were not produced by the principal pair. Parentage of 4.1% of the juveniles was completely outside the social group; these appear to result from early dispersal of juveniles rather than from brood parasitism, most likely resulting from breakup of nearby groups. Principal females mated outside of their group (2.3%), or with an auxiliary male (2.3%), in the remaining cases of parentage outside the principal pair. No matings were detected between close relatives (e.g. mother-son); matings detected between the principal female and an auxiliary male followed a typical replacement of the principal female by an unrelated immigrant female. Our finger-printing results indicate that: (1) current fitness benefits accruing to most auxiliaries do not exceed their likely reproductive success had they dispersed successfully to a breeding position; (2) nearly all wren mating is monogamous and (3) behavioral dominants (especially females) can monopolize breeding. Received: 23 September 1994/Accepted after revision: 10 June 1995     

Weight comparisons and litter size manipulation in Columbian ground squirrels (Spermophilus columbianus) show evidence of costs of reproduction

I studied reproductive costs in the female Columbian ground squirrel (Spermophilus columbianus) using individually marked animals. I compared weight changes during the active season and over winter, and mortality for females that did and did not wean young. Females raising young were heavier at emergence in that spring than unsuccessful ones. Females that did not raise young gained more weight during summer, were heavier than successful females at the time of entry into hibernation, and were heavier emerging from hibernation the following spring. Over-winter mortality was higher for females that reared young compared to reproductively unsuccessful females. A food supplementation experiment showed that energy-rich food can accelerate individuals’ weight gain. Interactions between litter size, birth weight, weight at emergence from the natal burrow, survival of young to yearling age, and maternal fitness were also studied. Litter sizes were experimentally manipulated to evaluate how females cope with costs of rearing one additional young. Birth weight of juveniles was positively correlated with survival to emergence from the natal burrow and with survival to yearling age. Partial litter loss was higher in experimentally enlarged litters than in either experimentally decreased or control litters. Total litter loss, survival of adult females or the probability of weaning young the following year were not affected by the litter size manipulation. Females appear to adjust the size of their litter before birth, and to some extent during lactation, to their ability to wean young. Received: 20 January 2000 / Received in revised form: 12 March 2000 / Accepted: 18 March 2000     

Do female bank voles (Clethrionomys glareolus) mate multiply to improve on previous mates?

Contrary to classical sexual selection theories, females of many taxa mate with multiple males during one reproductive cycle. In this study, we conducted an experiment on the “trade-up hypothesis”, which proposes that females remate if a subsequently encountered male is potentially superior to previous mates to maximize the genetic quality of their offspring. We presented bank vole females (Clethrionomys glareolus) sequentially with two males of known dominance rank in different orders, i.e., either first subordinate and second dominant, first dominant and second subordinate, or two males that were equal in dominance (high ranking) and observed their mating behavior. We found that 92% of the females mated multiply and did not base their remating decision on male social status. Therefore, polyandry cannot be explained by the “trade-up hypothesis” based on dominance rank in this species. However, we found that dominant males sired significantly more offspring than subordinate males. This varied according to mating order: dominant males sired more offspring when they were second than when they were first. Moreover, litter sizes were significantly smaller when the dominant male was first (smallest relative success of dominant males) compared to litter sizes when mating order was reversed or both males equal in status. Our results suggest that even though multimale mating includes males that are of poorer quality and thus potentially decreases the fitness of offspring, most of a female’s offspring are sired by dominant males. Whether this is due to cryptic female choice, sperm competition, or a combination of both, remains to be tested.