Who cares? Individual contributions to pup care by breeders vs non-breeders in the cooperatively breeding banded mongoose (<Emphasis Type="Italic">Mungos mungo</Emphasis>)

Knowledge of the investment rules adopted by breeders and non-breeders, and the factors that affect them, is essential to understanding cooperative breeding as part of a life-history tactic. Although the factors that affect relative contributions to care of young have been studied in some cooperative bird species, there is little data on mammals, making coherent generalisations within mammals and across taxa difficult. In this study, we investigate individual contributions to pup escorting, a strong predictor of offspring provisioning, in the banded mongoose (Mungos mungo), a cooperatively breeding mammal in which reproductive skew is low. Contributions by those under a year old (which virtually never breed) increased with age and body weight but were generally low. Among older age classes (yearlings and adults), individuals that had not bred in the current litter generally contributed less to escorting than those that had bred (with the exception of yearling males). In addition, females that did not breed reduced their investment if they were heavy presumably because such females are more likely to breed in the following event and benefit from saving resources for this. The generally greater contributions by breeders in banded mongooses contrast with the recent findings in meerkats (Suricata suricatta), another obligatorily cooperative mongoose with similar group size but wherein reproductive skew is high. Our results suggest that relative contributions by breeders vs non-breeders are not dependent on group size but on the ratio of breeders to carers and the probability that non-breeders will breed in the near future.     

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     

Paternity analysis shows experience,not age,enhances mating success in an aquatically mating pinniped,the Weddell seal (<Emphasis Type="Italic">Leptonychotes weddellii</Emphasis>)

For polygynous mammals with no paternal care, the number of offspring sired is often the sole measure of male reproductive success. The potential for polygyny is highest when resources or other environmental factors such as restricted breeding sites force females to aggregate. In these circumstances, males compete intensely for females and mating success may vary greatly among males, further intensifying selection for those traits that confer an advantage in reproduction. Hence, determinants of male success in competition for females are likely to be under strong sexual selection. Paternity analysis was used in conjunction with measures of age, site fidelity, and behavior during the breeding season to assess variance in male breeding success in Weddell seals (Leptonychotes weddellii) breeding at Turtle Rock, McMurdo Sound (77.727S, 166.85E) between 1997 and 2000. Paternity could be assigned to 177 pups at relaxed or 80% confidence level or 111 pups at strict or 95% confidence levels. Weddell seals at Turtle Rock show a modest degree of polygyny with the greatest number of pups sired by any individual male in a single season equalling 5 or ∼10% of the pups born. Over four consecutive years, most (89.2%) males sired at least one pup. In a generalized linear model (GLM), age and the age first seen at the study site as an adult were unrelated to mating success, but adult experience, either site-specific or elsewhere in McMurdo Sound, over the reproductive life span of males explained nearly 40% of variance in total mating success with 80% confidence and 24% of variance at 95% confidence. While learning where females are likely to be may enhance male reproductive success, aquatic mating reduces the ability of males to monopolize females, and thereby increases equity in mating success.     

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.     

Communal nursing in the evening bat,Nycticeius humeralis

Summary Nursing observations over two summers involving 76 lactating female evening bats, Nycticeius humeralis, and 128 pups in an attic in northern Missouri indicate that communal nursing occurs rarely until 2 weeks before weaning during which time over 18% of nursing bouts involve nondescendant offspring. The average relatedness among female pairs nursing non-descendant offspring, based on identity-by-descent estimates using allozyme data, was 0.04 (SE=0.12). Mitochondrial DNA d-loop sequence comparisons confirm that at most only 2 of 20 female pairs nursing non-descendant offspring came from the same matriline. Thus, females do not nurse matrilineal kin preferentially despite female natal philopatry. In addition, the average degree of relatedness within a colony (r=0.01, SE =0.03) is too low to provide any indirect benefits from communal nursing. Female error alone is insufficient to explain these observations because females tended to allow female nondescendant young to nurse but excluded nondescendant males, particularly when they had all-male litters. Furthermore, communal nursing bouts did not differ in duration from parental nursing bouts and involved 31 % of all banded females and 24% of all banded pups observed nursing. Communal nursing occurred most frequently when pups began hunting on their own and when lactating females attained their lowest average pre-fed body weight. Mortality during this period was higher for male than female pups, and relative weights implicate starvation as the cause. Time-lapse video records of four families of bats in captivity showed that the number of nursing bouts was proportional to daily weight change. I propose that these results are consistent with both immediate and delayed benefits accruing to females which experience variable hunting success. If a female with extra milk reduced her weight by dumping milk prior to her next foraging trip, she could obtain an immediate energetic benefit and maintain maximum milk production. By restricting such milk donations to nondescendant females she may also increase colony size and thereby enhance her future acquisition of information about foraging and roosting sites.     

Genetic reconstruction of breeding patterns in gundis (Rodentia: Ctenodactylidae)

Current mating system theory predicts that the number of females breeding in a group will depend on the number of females in the group and the accessibility of unrelated males, whereas the number of males breeding in a group will depend on the ability of males to control access to reproductive females. By combining information on group composition with genetic data, I determined whether breeding patterns in a rock-dwelling rodent, Ctenodactylus gundi, were concordant with these expectations. C. gundi breeding patterns varied from facultative monogamy to uni-male polygyny to multi-male polygyny. The number of reproductive individuals of each sex in a group increased with group size. Whereas communal breeding among related females tended to increase female reproductive success, males breeding in the same group were unrelated to other group members and seemed to compete over access to matings. While some males were assigned offspring from neighboring social groups, most group-living males successfully monopolized the reproduction of females within their group. There was no evidence that females had multiply sired litters, although some bred with alternative males in separate breeding attempts. Although numerous individuals were not assigned as parents or offspring, genetic information enabled me to determine that most unassigned individuals were philopatric group members, whereas only a few were unrelated immigrants into their current social group. Together, these results provide evidence that C. gundi social groups represent fairly distinct reproductive units whose breeding patterns are dependent on group size and composition in accordance with theoretical predictions. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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     

Harem choice and breeding experience of female southern elephant seals influence offspring survival

Female mammals can increase their lifetime fitness through modification of investment potential and by providing better rearing environments with improved breeding experience. We examined the relationships between reproductive fitness and the behavioural decisions that female southern elephant seals (Mirounga leonina) made during the breeding season. We examined whether mother age and breeding experience influenced reproductive success (measured as 1st-year survival probability), and whether there was a change in the choice of harem size with increasing age. Pups produced by young mothers had lower 1st-year survival probability than pups produced by older mothers. A significant increase in mean female mass with age required an analysis of both these effects on offspring survival. There was a significant positive effect of both female age and mass, and the interaction between the two, on 1st-year pup survival. The proportion of young mothers (<5 years old) decreased and the proportion of older mothers (>6 years old) increased with increasing harem size (harems surveyed from 1997 to 2001). Females chose larger harems in which to breed as they aged. Females demonstrated fidelity to breeding areas among successive breeding seasons, with older females displaying greater breeding-site fidelity than younger females. The mean number of previous breeding attempts per female within a harem (breeding experience) increased significantly with increasing harem size. Breeding females returned to breed later in the breeding season as they aged—we hypothesize that young, subordinate females gain a priority advantage by returning earlier. These results lend support to the hypothesis that there are fitness advantages, in terms of offspring survival, that are conferred to females that breed in successively larger harems with age. Potential mechanisms that select for females to improve their breeding conditions include improved mate selection and the avoidance of conspecific harassment in harems.Communicated by F. Trillmich     

Communal suckling in the cavy Galea musteloides

We quantified the extent of communal suckling in the cavy Galea musteloides. Six groups of animals were held in large indoor enclosures and suckling behavior was recorded over 113 h of observation. The groups contained 2–6 lactating females and 3–14 sucking pups. Due to the relative synchronization of births, 73% of the pups present in each group during lactation were non-offspring. Each of the 22 lactating females in the six groups suckled non-offspring in addition to her own offspring. On average, females suckled 86% of non-offspring present in their groups. Thus, 98% of all pups (n = 47) received milk from non-mothers. Although suckling frequencies were significantly higher for mothers with their own individual offspring than with non-offspring individuals, females invested more total time suckling all non-offspring than did suckling just their own; this was possible because for each mother many more non-offspring than offspring pups were present during lactation. Suckling bouts were significantly longer for mothers with their own individual offspring than with non-offspring individuals. The proportion of non-offspring suckling of mothers correlated negatively with the proportion of own young among the pups of a group. Non-offspring suckling did not affect future reproduction of females. Our observations demonstrate extensive practice of communal suckling in G. musteloides under laboratory conditions. Probably because all mothers of a group participated more or less equally in communal suckling behavior, the obvious cost of giving energetically expensive milk to non-offspring did not result in reduced (future) reproductive success. Potential benefits directly involved with communal suckling are unclear. More indirectly, communal suckling as well as birth synchrony might contribute to the formation of advantageous multi-litter kindergardens.     

Communal nesting, kinship, and maternal success in a social primate

Communal nesting, where several mothers regularly pool and cooperatively rear offspring, is unusual in mammals. This type of crèching behavior is especially rare among primates, with the notable exceptions of humans, some nocturnal strepsirrhines, and—as we show in this study—black-and-white ruffed lemurs (Varecia variegata). Here, we combine data on nesting behavior, genetic relatedness, and infant survivorship to describe variation in ruffed lemur infant care and to examine the potential benefits of ruffed lemur communal breeding. Reproductive events were rare, and females produced litters (synchronously) only once in 6 years of observation. We show that not all mothers participate in communal crèches, but those that did had greater maternal success; communal breeders spent more time feeding and their offspring were more likely to survive. Although cooperating mothers were often related, females also cooperated with non-kin, and those who shared infant care responsibilities had greater maternal success than mothers who did not participate. If there is indeed a causal link between maternal cooperation and reproductive success, this unusual behavior, like that of human communal rearing, may have evolved via some combination of kin selection and mutualism.     

Costs and benefits of communal rearing to female prairie voles (<Emphasis Type="Italic">Microtus ochrogaster</Emphasis>)

Communal rearing of offspring may help mothers maximize their investment in offspring at a reduced cost to their own bodily condition, thus maximizing their potential for reproductive success. The objective of this study was to quantify the costs and benefits of communal rearing to prairie vole (Microtus ochrogaster) pups and mothers. Mothers were assigned to one of three social units: solitary mothers, singularly breeding groups (i.e. one mother and one non-reproductive sister) and plurally breeding groups (i.e. two lactating sisters). For each type of social unit, some replicates were provided with food ad libitum, while others were provided with limited food. The body mass of focal mothers (i.e. the first mother to produce a litter) was a significant predictor of pup growth. Regardless of food availability, litters of focal mothers in plurally breeding groups gained more weight than litters reared by solitary mothers. Pups reared in singularly breeding groups were intermediate in weight gain, but did not gain significantly more weight than solitary offspring. There was no difference in the body mass of focal mothers from each type of social unit, regardless of food availability. Within plurally breeding groups, the weight gain of the two litters and body mass of focal and second mothers did not differ. However, focal mothers from plurally breeding groups nursed fewer pups than solitary mothers and also fewer pups than their nestmates when food was limited. Our results suggest that plural breeding results in greater fitness to mothers than solitary and singular breeding.Communicated by E. Korpimäki     

Synchrony in egg-laying and reproductive success of neighboring common murres,Uria aalge

Common murres (Uria aalge) are highly colonial; pairs often breed at the highest possible densities, in bodily contact with neighbors. At Bluff, a colony in western Alaska, we tested for synchrony in egg laying at various spatial scales and found little evidence for higher synchrony, either within study plots of 15–195 pairs, or within subplots containing several pairs, than among plots in a 5-year study. Egg laying of neighbors generally was more synchronous than expected based on overall frequency distributions in laying dates, however. Breeding success was positively correlated with the number of breeding neighbors and the number of neighbors tending eggs or nestlings at the time of egg laying. Breeding success of pairs with neighbors was positively related to the breeding success of neighbors. Pairs that produced eggs synchronously with at least one neighboring pair had higher success than those that began breeding either before or after their neighbors. Most reproductive failures at Bluff are due to accidental egg loss and predation on eggs by common ravens, Corvus corax, soon after laying. By occupying space where a raven might otherwise land and defending their own eggs, active breeding neighbors locally reduce the probability of egg predation. Active breeding neighbors also are less likely to flush and accidentally dislodge nearby eggs when disturbed than are nonbreeders. Murres breeding synchronously with neighbors have the highest assurance of the presence of active breeding neighbors both at the time of egg laying and throughout their reproductive attempts. Groups of neighboring murres can be considered small “selfish herds,” demonstrating by-product mutualism through their continued presence and defense of their own eggs and nestlings. Despite the advantages of breeding synchronously with neighbors, early breeding may often be favored, however. Received: 22 January 1996/Accepted after revision: 16 June 1996     

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.     

What do pairing patterns in common tern, Sterna hirundo, recruits reveal about the significance of sex and breeding experience?

In general, reproductive output in long-lived bird species increases in older compared to younger individuals. Therefore, experienced mates should be advantageous for first-time breeders. To examine requirements and consequences of experienced pair mates we investigated the first pair bonds of common terns, Sterna hirundo, recruiting to their natal colony. We found that male recruits were usually the same age as their mates, whereas female recruits were usually the younger member of the pair. In order to acquire experienced mates, it was necessary for males to arrive early in the year of recruitment. Mates with 2 or more years of breeding experience were only attainable by male recruits characterised by greater body mass and age. Female recruits arrived more than 1 week later than their experienced mates and significantly later in the season than male recruits paired with experienced females. In general, females first bred at a younger age than males, and neither the female recruits body mass nor their age was related to the level of experience of their first mate. These sex-specific differences in obtaining an experienced mate did not result in different levels of reproductive success between the sexes. Male and female recruits with mates with 2 or more years breeding experience benefited from having experienced mates: they had greater reproductive success. First-time breeders paired with mates with only 1 year of breeding experience did not differ from pairs where both members were breeding for the first time in terms of reproductive success, but clutches were larger. Our results illustrate not only different prerequisites for males and females, but also males need for experienced mates. Delayed male breeding (postponing breeding for another year), supposed to be a negative trait, and high body mass, supposed to be a trait of superior individual quality, can be combined in some individuals, improving reproductive success and showing that breeding common terns use a range of tactics to begin reproduction.Communicated by F. Trillmich     

Effectiveness of territorial polygyny and alternative mating strategies in northern fur seals, <Emphasis Type="Italic">Callorhinus ursinus</Emphasis>

We conducted a 6-year longitudinal behavioral and genetic investigation of a highly polygynous pinniped, the northern fur seal (Callorhinus ursinus), to determine the contribution of terrestrial polygyny to male fertilization success and to assess the occurrence of alternative mating strategies. Genetic samples from 37 adult males, 50 adult females, and 85 pups were collected and genotyped using five polymorphic microsatellite loci. Pup paternity was assigned using Cervus 2.0 at 99% confidence level. Paternity of 83 pups (98%) was assigned to terrestrial males who held territories or stayed temporarily in the study area during the breeding season when fertilization occurred. For 56 pups of which attendance records of their mothers were available, paternity of 45 pups (80%) was assigned to the associate males in whose territory their mothers stayed during the perioestrus period. In addition to defending breeding territories, territorial males have often been observed attempting to forcibly abduct adult females from adjacent territories (female stealing): We observed a total of 95 such cases, in which the stealers had significantly fewer females than the territorial males from whose territories they stole females. Our results indicate that terrestrial resource-defense polygyny is the major mating system in this species and that nonassociated paternity occurs mostly as a result of alternative mating strategies of less successful males. Male northern fur seals thus appear to adopt conditional alternative strategies that depend on their current social status to maximize their life-time reproductive success.     

The cost of success: reproductive effort in male southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>)

Reproductive effort is a key parameter of life history because it measures the resources allocated to reproduction at the expense of growth and maintenance. Male reproductive effort always had a minor role with respect to female effort both in the development of theories and in field research. Elephant seals are an ideal subject for reproductive effort studies because they fast during the breeding season, splitting the phase of energy acquisition from the phase of energy use for breeding. In this paper, we present results on male reproductive effort (weight loss estimated by photogrammetry) in southern elephant seals (Mirounga leonina), the most dimorphic and polygynous of all mammal species. We show that total reproductive effort increases with age, with no sign of late decrease or senescence. Male reproductive effort in this species depends mostly on behavioral factors, i.e., the success in competition with other males, and the intensity of interaction with females. A large effort results in large gains in both mating success and fertilizations. The large reproductive success that a few males are able to achieve come at a big cost in terms of energy expenditure, but this cost does not seem to affect the likelihood of survival to the following breeding season.     

Reproductive success of solitary and communal Philoponella oweni (Araneae: Uloboridae)

Summary Philoponella oweni (Uloboridae) is a facultatively communal orb-web spider found in arid regions in the southwestern United States. Within one habitat solitary adult females and communal groups of adult females coexist.This study compared the reproductive output (egg production) and reproductive success (production of live offspring) of solitary and communal females. There was no significant difference in the mean number of egg-cases produced by solitary and communal females, but communal females produced significantly more eggs per egg-case. However, communal females suffered significantly higher rates of egg-case parasitism from the pteromalid wasp Arachnopteromalus dasys. As a result, the mean number of liver offspring produced by solitary and communal females did not differ. Thus, both strategies led to roughly equal reproductive success. The advantages of aggregation of adults may be significant in the evolution of social behavior in spiders.     

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.     

The dynamics of brood desertion among communally breeding females in the treehopper, Publilia concava

In species that exhibit extended parental care, females sometimes lay eggs communally in order to redistribute the costs of offspring care. Communal egg-laying often involves redundancy in female effort, such that the number of females contributing to reproduction is more than is needed to provide adequate parental care. As a result, a subset of females will often abandon the communal brood, with the time of departure ranging from immediate abandonment after egg-laying (brood parasitism) to delayed abandonment with prolonged care (cooperative breeding). In this paper I approach the parental care dynamics of female-female broods as a desertion game similar to that of mate desertion in species with bi-parental care. I describe a field study of the insect Publilia concava (Hemiptera: Membracidae), a species that exhibits communal oviposition and a full range of egg guarding. This species exhibits full redundancy in female care, with no difference in survival between singly and doubly guarded broods. I find that double guarding is extremely rare in the population, with most communal broods having only one female guard. While this guard was usually the female that initiated the brood, these same females were more likely to abandon when secondary females arrived and when secondary females exhibited longer guarding durations. Paradoxically, the secondary females usually abandoned the broods they visited, resulting in up to 50% of broods with double abandonment. These unguarded broods suffered a 50% reduction in hatching success, reflecting an important risk for primary females that abandon egg masses to secondary females. Overall, P. concava exhibits desertion dynamics similar to mate desertion in vertebrates and it is likely that the theoretical work in this area will be useful for future work that addresses the allocation of parental care among communal breeders.     

Reproduction by subordinates in cooperatively breeding Arabian babblers is uncommon but predictable

In a genetic analysis of the mating system of cooperatively breeding Arabian babblers (Timalidae: Turdoides squamiceps), we identified which individuals in the population are breeding, and how reproductive success was distributed among group members with respect to their dominance rank, for both males and females. The population was characterized by an asymmetrical distribution of reproductive success; behaviorally dominant males produced 176 of 186 (95%) of the offspring in 44 social groups analyzed, and alpha females produced 185 of 186 (99.5%). We evaluated models of reproductive skew by examining genetic and demographic correlates of reproduction by␣subordinates. Subordinate (beta) males that sired young were more likely to be recent dispersers from their natal groups or members of newly formed groups than betas that did not reproduce. Breeding beta males had spent smaller proportions of their lives with the current alpha male and female as alphas than had beta males that did not sire young. One consequence of the linkage of dispersal with breeding in newly formed, nonnatal groups is that beta males that sired young had significantly lower genetic similarity to the alpha males in their groups (based on band-sharing coefficients using multilocus minisatellite DNA fingerprinting) than those that did not sire young. This pattern may occur generally in species in which group membership accrues both through nondispersal of young (forming groups of relatives) as well as through dispersal involving coalitions that sometimes include nonrelatives. Received: 22 July 1997 / Accepted after revision: 5 February 1998