Complete reproductive skew within white-browed sparrow weaver groups despite outbreeding opportunities for subordinates of both sexes

The distribution of reproductive success within societies is a key determinant of the outcomes of social evolution. Attempts to explain social diversity, therefore, require that we quantify reproductive skews and identify the mechanisms that generate them. Here, we address this priority using life history and genotypic data from >600 individuals in 40 wild groups of the cooperatively breeding white-browed sparrow weaver, Plocepasser mahali. We show that groups comprise up to six males and seven females, but within-group reproduction is completely monopolised by a single dominant male and female, while extra-group males sire 12–18 % of offspring. Strong within-group kin structure could frequently explain these monopolies, as subordinates had typically delayed dispersal from their natal groups and so frequently (1) lacked within-group outbreeding partners, and/or (2) stood to gain little from contesting dominant reproduction, being almost as related to the dominant’s young as they would have been to their own. Kin structure alone cannot account entirely for these monopolies, however, as they remained complete following the immigration of unrelated males and females. That subordinate females remain reproductively quiescent despite also showing comparable body condition to dominants, overlapping them substantially in age, and showing no evidence of elevated stress hormone levels raises the possibility that they exercise reproductive restraint due instead to a threat of action by dominants and/or deficits in offspring fitness that might arise if subordinates bred. Our findings highlight the complexity of the mechanisms that generate reproductive disparities in animal societies and the challenge of identifying them when skews are complete.     

Tug-of-war over reproduction in a cooperatively breeding cichlid

In group-living animals, dominants may suppress subordinate reproduction directly and indirectly, thereby skewing reproduction in their favour. In this study, we show experimentally that this ability (‘power’) is influenced by resource distribution and the body size difference between unrelated dominants and subordinates in the cichlid Neolamprologus pulcher. Reproduction was strongly skewed towards the dominant female, due to these females producing more and larger clutches and those clutches surviving egg eating better than those of subordinate females, but was not so when subordinates defended a patch. If breeding shelters were provided in two patches, subordinate females were more likely to exclusively defend a patch against the dominant female and breed, compared to when the same breeding resource was provided in one patch. Relatively large subordinate females were more likely to defend a patch and reproduce. Females also directly interfered with each other’s reproduction by eating the competitors’ eggs, at which dominants were more successful. Although dominant females benefited from subordinate females due to alloparental care and an increase in egg mass, they also showed costs due to reduced growth in the presence of subordinates. The results support the view that the dominant’s power to control subordinate reproduction determines reproductive partitioning, in agreement with the predictions from tug-of-war models of reproductive skew.     

Reproductive sharing and proximate factors mediating cooperative breeding in the African wild dog (Lycaon pictus)

Although dominant African wild dogs (Lycaon pictus) are generally believed to be the sole breeders within a pack, earlier behavioral and endocrine data suggest that reproduction could be shared with subordinates. We performed an extensive behavioral, demographic, and genetic evaluation of a wild dog population in South Africa to examine the level of such sharing and the proximate mechanisms influencing reproductive contributions of each sex. While a majority of pups were born to dominants because of a lack of subordinate potential breeders, we discovered a substantial portion of reproductive sharing between dominants and subordinates. Compared with alpha females that mated annually, subordinate beta females bred in 54.5% of years whereas thetas never bred. The three top-ranking males all sired pups (56.0%, 32.0%, and 12.0%, respectively) when three or more adult males were present. With only two pack males, alpha and beta individuals shared reproduction nearly equally (55.2% and 44.8%, respectively), and litters of mixed paternity were discovered on eight of 15 (53.3%) occasions. A skewed adult sex-ratio and frequent alpha mortalities for females and behavioral aggression in males allowed most individuals to attain dominant status in their lifetime, creating a constantly shifting social hierarchy. Genetic parentage results corresponded to reported hormone profiles, suggesting physiological suppression in some lower-ranked individuals of both sexes. Thus, a combination of demographic, behavioral, and hormonal proximate factors mediates reproductive partitioning in wild dogs. We conclude that reproductive sharing can be significant in this species, especially for males that have less robust suppressive mechanisms than females.     

Do stress hormones suppress helper reproduction in the cooperatively breeding red-cockaded woodpecker (<Emphasis Type="Italic">Picoides borealis</Emphasis>)?

In the cooperatively breeding red-cockaded woodpecker (Picoides borealis), male helpers are subordinate to male breeders and do not mate with females, even when unrelated to the breeding female within their group or through extra-group matings, yet exhibit reproductive hormone profiles similar to those of breeders. We investigated whether reproduction might be suppressed in helper males via high levels of the stress hormone corticosterone. We also examined effects of group size and season on corticosterone levels by comparing baseline and maximal plasma levels of corticosterone between helper males and breeding males, and among helper males and breeders of both sexes living in groups of different sizes throughout the reproductive cycle. We also measured plasma levels of estrogen, progesterone, and testosterone to examine other potential hormonal differences between helpers and breeders. Male status did not explain variation in any hormones; therefore, our data do not support the hypothesis that helper males are reproductively suppressed via corticosterone or the other hormones investigated. However, the presence of two or more helper males in a group tended to reduce baseline corticosterone in breeding and helper males, but not breeding females, suggesting that helper males reduce parental effort of other male group members. Seasonally, maximal corticosterone peaked during the nestling provisioning phase for breeding and helper males, but not breeder females, suggesting that males show an increased response to stressors posing a potential threat to survival of offspring.     

Dominance status influences female reproductive strategy in a territorial African rodent <Emphasis Type="Italic">Rhabdomys pumilio</Emphasis>

Dominance status influences the fitness of many mammals. Using African striped mice Rhabdomys pumilio, we tested whether (1) dominant females have greater reproductive success than subordinate females, (2) dominant females influence the reproductive output of subordinate females when they are housed in close proximity, (3) reproductive output of a female changes in response to the dominance status of her neighbours, and (4) whether prolonged association between individuals influences the variance in reproductive success between dominants and subordinates (i.e. the ‘dear enemy’ phenomenon). The size and mass of litters of dominants increased significantly when housed adjacent to subordinates than when housed apart. The litter size and mass of subordinates remained unchanged, although subordinates spent significantly more time with their pups when housed close to dominants than when housed apart; time spent with pups by dominants remained unchanged. Moreover, females modified their reproductive output and behaviour in relation to the dominance status of their neighbours. Following prolonged association, dominants still had greater reproductive success, but now, the time spent with pups decreased in subordinates. We suggest that dominants adopt a strategy to increase the reproductive value of their litter, whereas subordinates adopt a pup defence strategy. These strategies are flexible and are influenced by the dominance status and period of association between neighbours, so that females could maximize their fitness in response to varying social conditions.     

Subordinate males are more likely to help if unrelated to the breeding female in cooperatively breeding white-browed scrubwrens

Subordinates in communally breeding groups of birds usually help to provision nestlings, but in some species helping-at-the-nest is facultative. In species in which groups usually contain relatives, subordinates either always feed young or are more likely to do so when breeding dominants are close relatives, suggesting that benefits of helping collateral kin are important. In other species, adult group members are unrelated to each other and males may only feed young if they have gained paternity, showing that cooperation is related to the mating system. The white-browed scrubwren, Sericornis frontalis, is a communally breeding species in which most groups consist of a simple pair or a dominant pair with a subordinate male. Subordinate males either fed nestlings in a given nest at a rate comparable to the dominants, or did not feed them at all. Breeding groups usually formed through natal philopatry of males, so that about 80% of subordinates were closely related to one or both members of the dominant pair. However, because of death and dispersal, 54% of subordinates were unrelated to the resident female. Although subordinates with their mother fed nestlings in 48% of cases, they fed offspring in 75% of cases if their mother had been replaced by an unrelated female, suggesting that their decision to help is influenced by the opportunity to mate with the female. Supporting this conclusion, relatedness to the dominant male did not affect subordinate behaviour, and genetic studies showed that subordinates often gained paternity if unrelated to the female. Thus, paradoxically, provisioning nestlings is related to the opportunity for mating in a society in which there is natal philopatry and subordinates are usually related to one or both members of the dominant pair. Received: 25 January 1997 / Accepted after revision: 1 May 1997     

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.     

Delayed breeding in the cooperatively breeding Florida scrub-jay (Aphelocoma coerulescens): inhibition or the absence of stimulation?

To determine whether fundamental differences exist in the reproductive physiology of breeder and nonbreeder Florida scrub-jays (Aphelocoma coerulescens), we compared plasma levels of testosterone (T) and luteinizing hormone (LH) in males, and estradiol (E₂) and LH in females. Although male breeders had higher overall T and larger testes, nonbreeders’ T paralleled that of breeders, and their testes were more than an order of magnitude larger than regressed testes. Breeder and nonbreeder males had equivalent baseline LH, and equivalent changes in LH following a gonadotropin-releasing hormone (cGnRH-I) challenge. The T, LH and GnRH challenge data indicate that nonbreeder males have functional hypothalamo-pituitary-gonadal (HPG) axes. We found no hormonal evidence of inbreeding suppression in males: nonbreeders that did not live with their mothers and those that did had similar T. Male nonbreeders that were exposed to E₂-implanted females had higher T than did controls, suggesting that the lack of within-pair stimulation is a key factor in whether an individual delays breeding. Female nonbreeders had E₂ titres equal to or higher than breeders and neither basal LH nor LH following GnRH challenge differed by breeding status. Nonbreeders’ ovarian follicles were smaller than breeders’, but were larger than they would be during the non-breeding season. These data suggest that nonbreeders were primed for breeding and were simply waiting for an opportunity or a required stimulus. Female nonbreeders that lived in a territory with an unrelated male breeder had significantly higher E₂ than those that remained with their fathers. Similarly, nonbreeders that were captured away from their home territories had elevated E₂. However, nonbreeders that lived with their fathers had E₂ that was equivalent to breeding females, suggesting that inbreeding avoidance may not be the primary factor leading to delayed breeding in females. Received: 13 June 1995 /Accepted after revision: 27 April 1996     

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.     

Kin-preferential cooperation,dominance-dependent reproductive skew,and competition for mates in communally nesting female house mice

Little is known about the behavioural mechanisms facilitating kin-preferential communal breeding in wild house mice (Mus domesticus). We evaluated the effect of kinship and male availability on aggression, social structure and reproductive skew in groups of female mice freely interacting and reproducing in semi-natural indoor enclosures. Triplets of either sisters or non-sisters were established in enclosures provided with either one or three littermate males, which were unrelated and unfamiliar to the females. Sisters were more spatially associated and less aggressive than non-sisters, leading to higher incidences of communal breeding and reproduction. This is in agreement with theoretical considerations on kin selection in house mice. Reproductive success was highly skewed in favour of dominant females due to subordinate infertility or complete loss of first litters, which might have been caused by dominant females. In spite of this, subordinates only rarely dispersed from the enclosures, suggesting that perceived dispersal risk generally outweighed relatively reduced reproductive potentials. Aggression levels among females were significantly higher when one male was available, compared to when three males were available. We suggest that this might result from higher female-female competition for mates, due to the risk of missing fertilisation when synchronously oestrous females encounter limited numbers of males in a deme. Our results indicate that, first, communal nursing in house mice might have evolved to make the best out of a bad job rather than to enhance offspring fitness; and, second, that female-female mate-competition might play an important role in shaping female social structure in this polygynous mammal.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by F. Trillmich     

Rank-related partner choice in the fission–fusion society of the spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>)

When social partners vary in their relative value, individuals should theoretically initiate partnerships with conspecifics of the highest value. Here, we tested this prediction in a wild population of spotted hyenas (Crocuta crocuta). Crocuta live in complex, fission–fusion societies structured by dominance hierarchies in which individuals vary greatly in their value as social companions. Because patterns of association among Crocuta reflect social preferences, we calculated association indices (AIs) to examine how social rank influences intrasexual partner choice among unrelated adults of both sexes. The highest-ranking individuals were generally most gregarious in both sexes. Females associated most often with dominant and adjacent-ranking females. Females joined subgroups based on the presence of particular conspecifics such that subordinates joined focal females at higher rates than did dominants. Dominants benefit from associations with subordinates by enjoying priority of access to resources obtained and defended by multiple group members, but the benefits of these associations to subordinates are unknown. To investigate this, we tested three hypotheses suggesting how subordinates might benefit from rank-related partner choice among unrelated females. We found that subordinates who initiated group formation benefited by gaining social and feeding tolerance from dominants. However, rates at which dominants provided coalitionary support to subordinates did not vary with AIs. Overall, our data resemble those documenting patterns of association among cercopithecine primates. We consider our results in light of optimal reproductive skew theory, Seyfarth’s rank attractiveness model, and biological market theory. Our data are more consistent with the predictions of Seyfarth’s model and of biological market theory than with those of skew theory.     

Mutual tolerance or reproductive competition? Patterns of reproductive skew among male redfronted lemurs (<Emphasis Type="Italic">Eulemur fulvus rufus</Emphasis>)

The social organization of gregarious lemurs significantly deviates from predictions of the socioecological model, as they form small groups in which the number of males approximately equals the number of females. This study uses models of reproductive skew theory as a new approach to explain this unusual group composition, in particular the high number of males, in a representative of these lemurs, the redfronted lemur (Eulemur fulvus rufus). We tested two central predictions of “concession” models of reproductive skew theory, which assume that subordinates may be allowed limited reproduction by dominant group members as an incentive to remain in the group, thereby increasing the group’s overall productivity. Accordingly, relatives are predicted to receive less reproduction than non-relatives, and the overall amount of reproductive concessions given to subordinates is predicted to increase as the number of subordinates increases. In addition, we tested whether the number of females in a group, a variable not previously incorporated in reproductive skew theory, affected reproductive skew among males. Using microsatellite analyses of tissue DNA, we determined paternities of 49 offspring born into our study population in Kirindy forest (western Madagascar) since 1996 to determine patterns of male reproductive skew to test these predictions. Our analyses revealed remarkable reproductive skew, with 71% of all infants being sired by dominant males, but both predictions of reproductive skew models could not be supported. Instead, the number of females best predicted the apportionment of reproduction among the males in this species, suggesting that current reproductive skew models need to incorporate this factor to predict reproductive partitioning among male primates and perhaps other group-living mammals. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Both Peter M. Kappeler and Markus Port contributed equally to this paper.     

Genital morphology linked to social status in the bank vole (<Emphasis Type="Italic">Myodes glareolus</Emphasis>)

Since genital morphology can influence the outcome of post-copulatory sexual selection, differences in the genitalia of dominant and subordinate males could be a factor contributing to the fertilisation advantage of dominant males under sperm competition. Here we investigate for the first time if penile morphology differs according to male social status in a promiscuous mammal, the bank vole (Myodes glareolus). In this species, dominant males typically achieve higher reproductive success than subordinates in post-copulatory sexual selection, and male genital morphology is complex, including both a baculum (os penis) and penile spines. Our results show that despite no difference in body size associated with male social status, baculum width is significantly larger in dominant male bank voles than in subordinates. We also found evidence of positive allometry and a relatively high coefficient of phenotypic variation in the baculum width of male bank voles, consistent with an influence of sexual selection. By contrast, baculum length and three measures of penile spinosity did not differ according to male social status or show evidence of positive allometry. We conclude that dominant male bank voles may benefit from an enlarged baculum under sperm competition and/or cryptic female choice and that differences in penile morphology according to male social status might be important but as yet largely unexplored source of variation in male reproductive success.     

Sperm storage reflects within- and extra-pair mating opportunities in a cooperatively breeding bird

In passerine birds, storage and maturation of sperm takes place in the cloacal protuberance (CP), an external swelling of the reproductive organ. The considerable variation in CP size among species is presumed to be a consequence of varying levels of sperm-competition, but whether individual variation in CP size within a species also reflects sperm competition is not well established. Here, we study temporal variation in male CP size in relation to within-pair and extra-pair mating opportunities and cuckoldry risk in purple-crowned fairy-wrens Malurus coronatus. This is a socially monogamous cooperatively breeding passerine that can breed year-round and has low levels of extra-pair paternity (in 6?% of broods). We show that male CP size sharply increased a few weeks before, and rapidly regressed after his partner laid eggs, consistent with a cost of its maintenance and/or sperm production. Surprisingly, despite low levels of extra-pair paternity, CP size of non-breeding and pre-breeding males was positively correlated with the number of breeding females in the population, suggesting that CP size is sensitive to extra-pair mating opportunities. However, CP sizes do not seem to reflect cuckoldry risk: CP size of dominant males was unaffected by the presence of a subordinate that was unrelated to the dominant female, although those subordinates occasionally sire offspring, and had a larger CP than subordinates living with their mother. Our results suggest that, even in a species with low levels of extra-pair paternity, individual investment in sperm storage reflects both within-pair and, albeit to lesser extent, extra-pair mating opportunities.     

The lemur syndrome unresolved: extreme male reproductive skew in sifakas (<Emphasis Type="Italic">Propithecus verreauxi</Emphasis>), a sexually monomorphic primate with female dominance

The primates of Madagascar (Lemuriformes) are unusual among mammals in that polygynous species lack sexual dimorphism, and females dominate males socially in most species. Moreover, lemur groups are relatively small and characterized by even adult sex ratios despite the fact that one male should be able to exclude other males from the group. One hypothesis to explain this combination of behavioral, morphological, and demographic traits (the “lemur syndrome”) postulates that male–male competition is relaxed and, hence, variance in male reproductive success is low. Reproductive skew theory provides a framework for testing this and several related predictions about lemur social evolution. Specifically, low reproductive skew is also predicted if dominant males or adult females make reproductive concessions to subordinates or if the latter group successfully pursues alternative reproductive tactics. However, suitable data on paternity, demography, and behavior for a conclusive test of these predictions have not been available in the past. In this paper, we show that male reproductive success in ten groups of Verreaux’s sifakas (Propithecus verreauxi) was extremely skewed in favor of the dominant male over 9 years. Our genetic analyses also revealed that more than a third of all groups are effectively harem groups because only one male was unrelated to the resident female(s). In groups with two or more non-natal males, the dominant sired 91% of 33 infants. Together, males pursuing one of several alternative reproductive tactics, such as roaming among several groups or immigrating peacefully, sired only 11% of infants. Thus, female sifakas do not control group composition by offering reproductive opportunities to subordinate males as staying incentives, intrasexual selection is not relaxed, and dominant males prevail in a tug-of-war over subordinate males. Because male reproductive skew in sifakas is even more pronounced than in harem-living anthropoids studied to date, intrasexual selection is clearly not relaxed, and the lemur syndrome is more puzzling than ever.     

Female control of reproductive skew in cooperatively breeding brown jays (<Emphasis Type="Italic">Cyanocorax morio</Emphasis>)

Brown jays (Cyanocorax morio) are long-lived, social corvids that live in large, stable, territorial groups (mean = 10 individuals). In this study, I determined the distribution of reproductive success within groups using multi-locus DNA fingerprinting. Breeding females produced virtually all (99%) of the young within their own nests. Reproduction within groups was highly skewed towards a single primary female, although long term data indicate that secondary females (female breeders that were usually younger and subordinate to the primary female) were sometimes successful. The high reproductive skew observed for females was associated with primary female aggression. Successful reproduction by secondary females may have been due to parental facilitation or the inability of primary females to completely suppress secondary females. Multiple paternity occurred in 31–43% of broods and extra-group paternity occurred in a minimum of 22% of broods. Patterns of paternity also varied between years, since females often switched or included new genetic mates. Although male consorts of nesting females fathered relatively few offspring (20%), they still had a higher chance of fathering offspring than any other single group male. Reproduction was less skewed for males than females as a result of female mating patterns. Female reproductive patterns are consistent with some of the predictions and assumptions from optimal skew models, while male reproductive patterns are not. The factors affecting skew in species with complex social systems such as incomplete control by breeders over subordinate reproduction, female control of paternity, and resource inheritance have not been well incorporated into reproductive skew models.Communicated by: J. Dickinson     

The costs and benefits of territory sharing for the Caribbean coral reef fish,Scarus iserti

Summary Along the Caribbean coast of Panama, groups of unrelated female striped parrotfish, Scarus iserti, co-defend a common feeding territory. Field manipulations of group size and composition were performed to examine the benefits and costs accrued by dominant fish within these shared territories. Dominant fish benefit from the presence of relatively large subordinates because they share in the defense of the territory. Removals of these fish caused increases in defense time and decreases in feeding time for dominant group members. Dominants benefit from the presence of small subordinates because they increase the foraging efficiency of dominants. Removals of smaller subordinates caused reductions in the feeding time of dominant fish, although no changes in defense time occurred. Concurrently, dominant fish reduce costs of resource depletion by displacing subordinate group members from good food patches. Dominance interactions within a group reduce the amount of time subordinates spend feeding (subordinate individuals fed at higher rates following the removal of a dominant) and limit a subordinate's access to high quality resources. This combination of benefits and reduced costs ultimately contributes to the economic defensibility of a striped parrotfish territory and has led to the evolution of group territorial behavior in the absence of kin selection and cooperative parental care.     

Fitness consequences of dispersal decisions for male mountain gorillas (Gorilla beringei beringei)

The fitness consequences of dispersal decisions are difficult to quantify, especially for long-lived species with complex social systems. To calculate those consequences for male mountain gorillas, from the perspective of both subordinate and dominant males, we used behavioral and demographic data obtained over 30 years from the Virunga Volcano population to develop an agent-based model that simulates the life history events, social structure, and population dynamics of the species. The model included variables for birth rates, mortality rates, dispersal patterns, and reproductive skew. The model predicted an average lifetime reproductive success (LRS) of 3.2 for philopatric males (followers) and 1.6 for emigrants. The benefits of philopatry were most sensitive to opportunities for social queuing and to female transfer preferences, but philopatry remained the best strategy over a wide range of group conditions and hypothetical simulations. The average LRS for dominant males was 4.5 when a subordinate stayed and 4.6 when the subordinate emigrated. The dispersal decision of the subordinate male had little impact on the fitness of the dominant male because it came relatively late in the dominant males reproductive life span, and it changed his group composition only incrementally. The fitness consequences for the dominant male were most sensitive to the degree of reproductive skew. Since subordinates suffer a fitness loss when they leave a group, they should accept whatever reproductive restraint is needed to avoid eviction, and the dominant male does not need to offer concessions for them to stay. The dominant male may offer reproductive concessions for other reasons, such as peace incentives or to confuse paternity, or he may not have complete control of reproduction within his group.Communicated by D. Watts     

Why breed communally? Factors affecting fecundity in a communal breeding mammal: the banded mongoose (<Emphasis Type="Italic">Mungos mungo</Emphasis>)

Fecundity is an important component of fitness. In cooperatively breeding species, studies aimed at understanding the factors that affect fecundity have largely been restricted to species that exhibit high reproductive skew, where reproduction is monopolised by a few individuals. In such species, dominant suppression and inbreeding avoidance are the principal explanations for low fecundity in subordinate females. In this paper, we evaluate the relative effects of individual, social, and environmental factors on female fecundity in a low skew cooperative breeding mammal: the banded mongoose (Mungos mungo). Most females (80%) conceived in each breeding event, and most pregnant females (93%) carried their litter to term. The principal determinants of a females fecundity were intrinsic qualities, particularly age and body size. However, there was no evidence of dominant suppression of subordinate reproduction or inbreeding avoidance. Similarly, there was little indication that social or environmental factors influence fecundity. We suggest that in the banded mongoose, the apparent lack of costs to inbreeding, and the absence of dominant female suppression of reproduction in other females result in low reproductive skew. Indeed, in banded mongooses, like lions (Felis leo), multiple breeding may be a consequence of benefits to rearing young communally.Communicated by F. Trillmich     

Rank-dependent hoarding effort in willow tits (<Emphasis Type="Italic">Parus montanus</Emphasis>): a test of theoretical predictions

Many birds and mammals store energy as hoarded food supplies. A supply of stored food is beneficial during periods when food is scarce, but building up and managing such a supply also entails costs. The optimal number of caches will be reached when the net benefit is at its maximum. If dominants can steal more stored food from subordinates than the other way around, the optimum will differ between these categories. A previous theoretical model of hoarding in groups with dominant and subordinate members produced three testable predictions: (1) hoarders should store more food as anticipated future conditions get worse; (2) subordinate flock members should store more food than dominants; and (3) dominants should increase hoarding relatively more than subordinates as conditions get worse. Here we present a field experiment on willow tits (Parus montanus) designed to test these predictions. We found support for all three. Hoarding increased as conditions got worse, subordinates stored at a higher rate than dominants, and dominants increased their hoarding effort relatively more than subordinates as conditions worsened. These results support the incorporation of information on dominance and food availability into models predicting food storage behaviour.Communciated by J. Dickinson