Genetic population structure and sociogenetic organisation in Formica truncorum (Hymenoptera; Formicidae)

Summary The genetic population structure and the sociogenetic organization of the red wood ant Formica truncorum were compared in two populations with monogynous colonies and two populations with polygynous colonies. The genetic population structure was analysed by measuring allele frequency differences among local subsets of the main study populations. The analysis of sociogenetic organisation included estimates of nestmate queen and nestmate worker relatedness, effective number of queens, effective number of matings per queen, relatedness among male mates of nestmate queens and relatedness between queens and their male mates. The monogynous populations showed no differentiation between subpopulations, whereas there were significant allele frequency differences among the subpopulations in the polygynous population. Workers, queens and males showed the same genetical population structure. The relatedness among nestmate workers and among nestmate queens was identical in the polygynous societies. In three of the four populations there was a significant heterozygote excess among queens. The queens were related to their male mates in the polygynous population analysed, but not in the monogynous ones. The data suggest limited dispersal and partial intranidal mating in the populations with polygynous colonies and outbreeding in the populations having monogynous colonies. Polyandry was common in both population types; about 50% of the females had mated at least twice. The males contributed unequally to the progeny, one male fathering on average 75% of the offspring with double mating and 45–80% with three or more matings. Correspondence to: L. Sundström     

Variation in colony structure in the subterranean termite Reticulitermes flavipes

The genetic organization of colonies of the subterranean termite Reticulitermes flavipes in two subpopulations in Massachusetts was explored using five polymorphic allozymes and double-strand conformation polymorphism (DSCP) analysis of the mitochondrial control region. Empirically obtained estimates of worker relatedness and F-statistics were compared with values generated by computer simulations of breeding schemes to make inferences about colony organization. In one study site (G), worker genotypes indicated the presence of a mixture of colonies headed by monogamous outbred primary reproductives and colonies headed by inbreeding neotenic reproductives, both colony types having limited spatial ranges. A second site (S) was dominated by several large colonies with low relatedness among nestmates. Mixed DSCP haplotypes in three colonies indicated that nestmates had descended from two or three unrelated female reproductives. Computer simulations of breeding schemes suggested that positive colony inbreeding coefficients at site S resulted from either commingling of workers from different nests or different colonies. Such an exchange of workers between nests corresponds to the multiple-site nesting lifetype of many subterranean termites and resembles colony structure in polycalic Formica ants. Our study demonstrates considerable variation in R. flavipes colony structure over a small spatial scale, including colonies headed by monogamous outbred primary reproductives, colonies containing multiple inbred neotenic reproductives and large polydomous colonies containing the progeny of two or more unrelated queens, and suggests that the number of reproductives and nestmate relatedness change with colony age and size.     

Intranest relatedness and nestmate recognition in the meadow ant Formica pratensis (R.)

The impact of intranest relatedness on nestmate recognition was tested in a population of polydomous and monodomous nests of the mound-building ant Formica pratensis. Nestmate recognition was evaluated by testing aggression levels between 37 pairs of nests (n=206 tests). Workers from donor colonies were placed on the mounds of recipient nests to score aggressive interactions among workers. A total of 555 workers from 27 nests were genotyped using four DNA microsatellites. The genetic and spatial distances of nests were positively correlated, indicating budding and/or fissioning as spread mechanisms. Monodomous and polydomous nests did not show different aggression levels. Aggression behavior between nests was positively correlated with both spatial distance and intranest relatedness of recipient colonies, but not with genetic distance or intranest relatedness of donor colonies. Multiple regression analysis revealed a stronger effect of spatial distance than of genetics on aggression behavior in this study, indicating that the relative importance of environment and genetics can be variable in F. pratensis. Nevertheless, the positive regression between intranest relatedness of recipient colonies and aggression in the multiple analysis supports earlier results that nestmate recognition is genetically influenced in F. pratensis and further indicates that foreign label rejection most likely explains our data.     

Ecology of<Emphasis Type="Italic"> Leptothorax</Emphasis> ants: impact of food,nest sites,and social parasites

In a long-term field manipulation, we demonstrate strong reactions of Leptothorax longispinosus ant colonies to food- and nest-site supplementation. Demographic and genetic responses varied over small geographic scales, and the two ecological factors interacted with the presence of the social parasite Protomognathus americanus. We conducted a 2×2 experiment in three blocks and found that the blocks, which were less than 100 m apart, reacted very differently to the treatments. Blocks differed in degree of polygyny, intranest relatedness, colony size, productivity, and sexual investment. Furthermore, these differences were associated with the presence of slave-making ants and the local availability of nest sites. Nest-site supplementation had a strong effect only in the site with the highest prevalence of social parasites, influencing there the density and investment patterns of colonies. L. longispinosus ants in the least parasitized area were strongly affected by both food- and nest-site supplementation. There, food supplementation led to a decrease in the number of queens per colony and consequently to an increase in intranest relatedness, while colonies in nest-site-supplemented areas invested fewer resources in males and produced a female-biased allocation ratio. By contrast, in a third block with a very low intracolonial relatedness, food supplementation induced an absolute and relative higher investment in males. We conclude that ecological factors influencing social organization in insect societies cannot be studied in isolation, because the interactions among factors produce far richer responses than any one variable.Communicated by L. Sundström     

Serial polygyny and colony genetic structure in the monogynous queenless ant Diacamma cyaneiventre

Serial polygyny, defined as the temporal succession of several reproductive females in a colony, occurs in some monogynous social insects and has so far attracted little attention. Diacamma cyaneiventre is a queenless ponerine ant found in the south of India. Colonies are headed by one singly mated worker, the gamergate. After the death of the gamergate or her absence following colony fission, the gamergate is replaced by a newly eclosed nestmate worker. After a replacement, colonies go through short-lived periods in which two matrilines of sisters co-occur. This is a situation which can be described as serial polygyny. To measure the consequences of serial polygyny, a genetic analysis was performed on 449 workers from 46 colonies of D. cyaneiventre using five microsatellite loci. The presence of more than one matriline among workers of the same nest was detected in 19% of colonies, indicating a recent change of gamergate. The average genetic relatedness among nestmate workers was 0.751 and did not significantly differ from the theoretical expectation under strict monogyny and monandry (0.75). A simple analytical model of the temporal dynamics of serial polygyny was developed in order to interpret these results. We show that the rate of gamergate turnover relative to the rate of worker turnover is the crucial parameter determining the level of serial polygyny and its effect on the genetic structure of colonies. This parameter, estimated from our data, confirms that serial polygyny occurs in D. cyaneiventre but is not strong enough to influence significantly the average genetic relatedness among workers.     

Mating and reproduction in the wasp Vespula germanica

We used polymorphic microsatellite markers to study patterns of queen and worker reproduction in annual nests of the wasp Vespula germanica in its introduced range in Australia. We found that queens were typically polyandrous (at least 85.4% mated multiply), with the minimum number of male mates ranging from 1 to 7. Calculations based on nestmate worker relatedness (r=0.46) yielded an estimate of effective queen mating frequency of 2.35. Queens were unrelated to their mates (r=-0.01), indicating that mating occurred at random within Australian V. germanica populations. In addition, the distribution of the minimum number of male mates of queens followed a Poisson distribution. This result suggested that the probability of a queen remating was not affected by previous copulations. We also discovered that mates of polyandrous queens contributed unequally to progeny production leading to significant male reproductive skew within nests. Analyses of nestmate male genotypes revealed that queens usually produced most or all males. However, workers were responsible for the production of many males in a few nests, and, in contrast to theoretical expectations, two of these nests were apparently queenright.     

Sociogenetic organization of the red ant Myrmica rubra

Knowledge of the sociogenetic organization determining the kin structure of social insect colonies is the basis for understanding the evolution of insect sociality. Kin structure is determined by the number and relatedness of queens and males reproducing in the colonies, and partitioning of reproduction among them. This study shows extreme flexibility in these traits in the facultatively polygynous red ant Myrmica rubra. Relatedness among worker nestmates varied from 0 to 0.82. The most important reason for this variation was the extensive variation in the queen number among populations. Most populations were moderately or highly polygynous resulting in low relatedness among worker nestmates, but effectively monogynous populations were also found. Polygynous populations also often tend to be polydomous, which is another reason for low relatedness. Coexisting queens were positively related in two populations out of five and relatedness was usually similar among workers in the same colonies. Due to the polydomous colony organization and short life span of queens, it was not possible to conclusively determine the importance of unequal reproduction among coexisting queens, but it did not seem to be important in determining the relatedness among worker nestmates. The estimates of the mating frequency by queens remained ambiguous, which may be due to variation among populations. In some populations relatedness among worker nestmates was high, suggesting monogyny and single mating by queens, but in single-queen laboratory nests relatedness among the worker offspring was lower, suggesting that multiple mating was common. The data on males were sparse, but indicated sperm precedence and no relatedness among males breeding in the same colony. A comparison of social organizations and habitat requirements of M. rubra and closely related M. ruginodis suggested that habitat longevity and patchiness may be important ecological factors promoting polygyny in Myrmica. Received: 15 May 1995/Accepted after revision: 17 October 1995     

Plasticity of social organization in a forest ant species

We manipulated availability of food and nesting sites in one population of the forest ant Myrmica punctiventris. The manipulations produced significant changes in relatedness structure, reproductive allocation, and response to hierarchical selection. Food availability appeared to have a consistently stronger influence on these aspects of social organization than did availability of nesting sites. We interpret our experimental results in light of observed differences between populations, and discuss implications for kin selection dynamics. Received: 30 July 1998 / Accepted after revision: 31 October 1998     

The genetic population structure of the gray mouse lemur (<Emphasis Type="Italic">Microcebus murinus), </Emphasis>a basal primate from Madagascar

The genetic structure of a population is closely connected to fundamental evolutionary processes and aspects of social behavior. Information on genetic structure is therefore instrumental for the interpretation of social behavior and evolutionary reconstructions of social systems. Gray mouse lemurs (Microcebus murinus) are basal primates endemic to Madagascar whose social organization is characterized by solitary foraging at night and communal resting during the day. Conflicting reports about population structure based on behavioral observations led us to examine the genetic structure of one population in detail in order to: (1) identify natural genetic units in this solitary primate, and (2) to test the assumption of current models of primate social evolution that solitary primates are organized in matrilines. DNA was extracted from tissue samples of 85 individuals from Kirindy forest to determine their variability at a 530 bp fragment of the mitochondrial D-loop and at six microsatellite loci. We found that this population was characterized by a great general diversity among mtDNA haplotypes, a pronounced sex difference in mtDNA haplotype diversity and spatial clustering of females with a particular haplotype, but low average relatedness among members of haplotype clusters. Specifically, we identified 13 different haplotypes, which were unevenly distributed among individuals. About 80% of all individuals, most of which were females or juvenile males, shared a single haplotype. Rare haplotypes were almost exclusively represented by single adult males, who apparently migrated into this population. One other haplotype was represented by a small group of females living at one edge of the study area. Microsatellite analysis revealed above-average relatedness among females with overlapping home ranges, as well as no signs of inbreeding, implying that male dispersal results in high levels of gene flow among matrilineal groups. We conclude that gray mouse lemur populations are hierarchically organized in small family units of closely related females that form stable sleeping groups, several of which are connected through a common mtDNA haplotype and form spatially distinct clusters. The presence of such matrilines supports a basic assumption of current models of primate social evolution.     

Seasonal nestmate recognition in the ant Formica exsecta

Under favorable conditions, the mound-building ant Formica exsecta may form polydomous colonies and can establish large nest aggregations. The lack of worker aggression towards nonnestmate conspecifics is a typical behavioral feature in such social organization, allowing for a free flux of individuals among nests. However, this mutual worker toleration may vary over the seasons and on spatial scales. We studied spatio-temporal variation of worker–worker aggression within and among nests of a polydomous F. exsecta population. In addition, we determined inter- and intracolony genetic relatedness by microsatellite DNA genotyping and assessed its effect on nestmate recognition. We found significant differences in the frequency of worker exchange among nests between spring, summer, and autumn. Moreover, we found significant seasonal variation in the level of aggression among workers of different nests. Aggression levels significantly correlated with spatial distance between nests in spring, but neither in summer nor in autumn. Multiple regression analysis revealed a stronger effect of spatial distances rather than genetic relatedness on aggressive behavior. Because nestmate discrimination disappeared over the season, the higher aggression in spring is most plausibly explained by cue intermixing during hibernation.     

Mean colony relatedness is a poor predictor of colony structure and female philopatry in the communally breeding Bechstein's bat (Myotis bechsteinii)

In order to understand why animals are social and how group members interact with each other it is important to know their relatedness. However, few studies have investigated the genealogy in complete social groups of free-living animals with low reproductive skew. This holds particularly true for bats. Although almost all bat species are social, their sociobiology is not well understood. Because they are volant, nocturnal and have a rather cryptic life-style, bats are difficult to observe in the wild. Furthermore females are generally gregarious making genetic parent-offspring assignment a challenging task. We used genetic markers in combination with knowledge about age and colony membership of individually marked bats to construct pedigrees in completely sampled maternity colonies of Bechstein's bats (Myotis bechsteinii). Despite considerable fluctuations in population size, no immigration occurred over 5 years in four colonies living in close proximity. Additionally, confrontation tests showed that females of one maternity colony were able to detect and attempted to prevent the intrusion of foreign females into a roost they occupy. Although colonies were absolutely closed, and 75% of the colony members lived together with close relatives (rS=0.25), mean colony relatedness was nearly zero (0.02). Average relatedness therefore is a poor estimator for the potential of kin selection in Bechstein's bat colonies and may be misleading when attempting to understand the social structure of animals living in groups where many members breed. Based on our results we discuss the potential adaptive value of living in closed societies with low reproductive skew.     

Queen primer pheromone affects conspecific fire ant (<Emphasis Type="Italic">Solenopsis invicta</Emphasis>) aggression

Monogyne fire ant, Solenopsis invicta, colony workers are territorial and are aggressive toward members of other fire ant colonies. In contrast, polygyne colony workers are not aggressive toward non-nestmates, presumably due to broader exposure to heritable and environmentally derived nestmate recognition cues (broad template). Workers from both monogyne and polygyne fire ant colonies execute newly mated queens after mating flights. We discovered that monogyne and polygyne queens have a remarkable effect on conspecific recognition. After removal of their colony queen, monogyne worker aggression toward non-nestmate conspecifics quickly drops to merely investigative levels; however, heterospecific recognition/aggression remains high. Queenless monogyne or polygyne worker groups were also not aggressive toward newly mated queens. Queenless worker groups of both forms that adopted a monogyne-derived newly mated queen became aggressive toward non-nestmate workers and newly mated queens. We propose that the powerful effect of fire ant queens on conspecific nestmate recognition is caused by a queen-produced recognition primer pheromone that increases the sensitivity of workers to subtle quantitative differences in nestmate recognition cues. This primer pheromone prevents the adoption of newly mated queens (regulation of reproductive competition) in S. invicta and when absent allows queenless workers to adopt a new queen readily. This extraordinary discovery has broad implications regarding monogyne and polygyne colony and population dynamics.     

Role of the nutritive value of sediment in regulating population dynamics of the deposit-feeding polychaete Streblospio shrubsolii

The importance of changes in nutritive value of the sediment in regulating the dynamics of the small deposit-feeding polychaete Streblospio shrubsolii (Buchanam, 1890) was examined. First, fluctuations in the population dynamics of S. shrubsolii and in the nutritive content of the sediment were studied on a seasonal scale. Subsequently, a manipulative field experiment was used to test the hypothesis that, in summer, availability of food regulates aspects of the population dynamics of S. shrubsolii. In summer 1997, sharp increases in protein concentrations and in the number of individuals of S. shrubsolii were found compared to all the other seasons. Densities of S. shrubsolii also showed an increase in the subsequent season. Newly recruited juveniles (19-31 setigers) were abundant in summer and autumn 1997, whereas the number of juveniles (32-42 setigers) increased in autumn 1997. Large numbers of adults were found in summer 1997. Significant negative correlations occurred in summer 1997 between the concentrations of proteins and the total number of individuals, the number of newly recruited juveniles and the density of adults. No differences in densities of juveniles, adults or total number of individuals were found among plots before starting the experiment. After 20 days of experimental enrichment, proteins showed a tendency to increase. After 40 days of enrichment, the total number of individuals was significantly greater in the enriched plots. Large numbers of newly recruited larvae and postlarval adults were found in the enriched plots. The experiment corroborated the model that the growth of population was limited by the availability of food in summer. Thus, in this season, patches of food could have a fundamental role in determining the population dynamics of small deposit-feeders under natural conditions and over short time scales.     

Experimental conversion of colony social organization by manipulation of worker genotype composition in fire ants (<Emphasis Type="Italic">Solenopsis invicta</Emphasis>)

Previous studies have shown that colony social organization in Solenopsis invicta is under strong genetic control. Colonies containing some proportion of workers with the Bb or bb genotypes at the gene Gp-9 display polygyne social organization (multiple reproductive queens per colony), whereas colonies with only BB workers express monogyne organization (single reproductive queen per colony). The hypothesis that the presence of workers bearing the b allele confers the polygyne social phenotype on a colony leads to the prediction that social organization can be manipulated by experimentally altering frequencies of adult workers bearing this allele. We did this by replacing queens in colonies of each social form with single queens of the alternate form, which differ in Gp-9 genotype. As worker Gp-9 genotype compositions changed, experimental colonies switched to the alternate social organization. These switches occurred when frequencies of workers with the b allele passed an identifiable threshold, such that colonies with fewer than 5% such workers behaved like monogyne colonies and those with more than 10% behaved like polygyne colonies. Our data thus confirm the prediction that colony social organization in this ant can be altered by manipulating adult worker genotype compositions, and thereby support the hypothesis that the expression of polygyny requires the presence of adult workers bearing the b allele at Gp-9.     

Invasive species impacts on ecosystem structure and function: A comparison of Oneida Lake, New York, USA, before and after zebra mussel invasion

Exotic species invasion is widely considered to affect ecosystem structure and function. Yet, few contemporary approaches can assess the effects of exotic species invasion at such an inclusive level. Our research presents one of the first attempts to examine the effects of an exotic species at the ecosystem level in a quantifiable manner. We used ecological network analysis (ENA) and a social network analysis (SNA) method called cohesion analysis to examine the effect of zebra mussel (Dreissena polymorpha) invasion on the Oneida Lake, New York, USA, food web. We used ENA to quantify ecosystem function through an analysis of food web carbon transfer that explicitly incorporated flow over all food web paths (direct and indirect). The cohesion analysis assessed ecosystem structure through an organization of food web members into subgroups of strongly interacting predators and prey. Our analysis detected effects of zebra mussel invasion throughout the entire Oneida Lake food web, including changes in trophic flow efficiency (i.e., carbon flow among trophic levels) and alterations of food web organization (i.e., paths of carbon flow) and ecosystem activity (i.e., total carbon flow). ENA indicated that zebra mussels altered food web function by shunting carbon from pelagic to benthic pathways, increasing dissipative flow loss, and decreasing ecosystem activity. SNA revealed the strength of zebra mussel perturbation as evidenced by a reorganization of food web subgroup structure, with a decrease in importance of pelagic pathways, a concomitant rise of benthic pathways, and a reorganization of interactions between top predator fish. Together, these analyses allowed for a holistic understanding of the effects of zebra mussel invasion on the Oneida Lake food web.     

Queen–worker–brood coadaptation rather than conflict may drive colony resource allocation in the ant Temnothorax curvispinosus

Conflicts of interest among genetically heterogeneous nestmates in social insect colonies have been emphasized as driving colony resource allocation. However, potential intracolonial conflicts may not actually be realized so that resource allocation could be shaped primarily by among-colony selection that maximizes colony productivity. To elucidate the causal basis of patterns of resource allocation, I experimentally manipulated three fundamental aspects of colony social structure (relatedness among workers, relatedness among larvae, and queen presence) in the ant Temnothorax curvispinosus and measured effects on colony resource allocation to new workers, gynes, and males. The experimental manipulations had widespread effects on patterns of colony resource allocation, but there was little evidence for realized conflicts over the sex ratio and caste ratio. Decreasing nestmate relatedness caused decreased colony productivity, suggesting that more closely related nestmates have more favorably interacting phenotypes. Together, these results suggest that resource allocation in T. curvispinosus may be shaped more by among-colony selection than intracolonial conflict, leading to queen–worker–brood coadaptation.     

Morphological, reproductive, and genetic variability among three populations of Crucibulum quiriquinae (Gastropoda: Calyptraeidae) in northern Chile

Crucibulum quiriquinae (Lesson, 1830) is the only species of Crucibulum currently recognized in northern Chile. Recent analysis of three Crucibulum populations obtained in northern Chile demonstrates the existence of morphological, genetic, and reproductive differences among populations. Two populations present in Bahía Tongoy (30°15'S), one inhabiting the shells of the snail Turritella cingulata and the other shells of the pectinid Argopecten purpuratus, showed morphological differences. However, both had planktonic larval development and show low genetic divergence (D=0.002). A third population from Bahía La Herradura (29°58'S), which also inhabits the shells of T. cingulata, did not show morphological differences compared with its counterpart from Bahía Tongoy. However the Bahía La Herradura population had intracapsular development and metamorphosis, and a larger genetic distance (D=0.06) from both Tongoy populations. The results of the reproductive and genetic analyses strongly suggested that the two Tongoy populations, although showing morphological differences, are biologically the same species, Crucibulum quiriquinae, whereas the La Herradura population is a new species.     

Cryptic species of Clavelina (Ascidiacea) in two different habitats: harbours and rocky littoral zones in the northwestern Mediterranean

Marinas and harbours provide ideal sites for the study of population genetics of marine invertebrates with restricted dispersal capabilities. They combine a confinement effect, particular ecological conditions (pollution, turbidity), and the possibility of high gene flow through ship-borne propagules, which greatly increases the natural dispersal capability of sexual and asexual propagules in many species with short-lived larvae. We studied the genetic structure of populations of the ascidian Clavelina lepadiformis living inside and outside harbours in the north-western Mediterranean. A 500-bp segment of the cytochrome c oxidase subunit I (COI) mitochondrial gene was sequenced in three populations from inside harbours (interior form) and in three populations from the rocky littoral (exterior form). Two congeneric Mediterranean species, Clavelina sp. and C. dellavallei, were used for comparison. We found that the interior and exterior forms of C. lepadiformis belong to two distinct clades, with a genetic divergence of 5%. Gene-flow values among these forms were insignificant. The lack of gene flow and the genetic divergence suggest that the interior and exterior forms of C. lepadiformis are in fact cryptic species rather than differentiated populations of the same species. Levels of gene flow were higher among interior habitats than among exterior habitats, a pattern likely maintained by genetic exchange through ships. We discuss the possible origins of the present-day distribution of these cryptic species. We contend that the study of species living both inside and outside these particular habitats will reveal more instances of genetic discontinuities allowing local adaptations.     

Reproductive conflicts and egg discrimination in a socially polymorphic ant

The ability to discriminate against competitors shapes cooperation and conflicts in all forms of social life. In insect societies, workers may detect and destroy eggs laid by other workers or by foreign queens, which can contribute to regulate reproductive conflicts among workers and queens. Variation in colony kin structure affects the magnitude of these conflicts and the diversity of cues used for discrimination, but the impact of the number of queens per colony on the ability of workers to discriminate between eggs of diverse origin has so far not been investigated. Here, we examined whether workers from the socially polymorphic ant Formica selysi distinguished eggs laid by nestmate workers from eggs laid by nestmate queens, as well as eggs laid by foreign queens from eggs laid by nestmate queens. Workers from single- and multiple-queen colonies discriminated worker-laid from queen-laid eggs, and eliminated the former. This suggests that workers collectively police each other in order to limit the colony-level costs of worker reproduction and not because of relatedness differences towards queens’ and workers’ sons. Workers from single-queen colonies discriminated eggs laid by foreign queens of the same social structure from eggs laid by nestmate queens. In contrast, workers from multiple-queen colonies did not make this distinction, possibly because cues on workers or eggs are more diverse. Overall, these data indicate that the ability of F. selysi workers to discriminate eggs is sufficient to restrain worker reproduction but does not permit discrimination between matrilines in multiple-queen colonies.     

Extended family structure in the ant Formica paralugubris: the role of the breeding system

In populations of various ant species, many queens reproduce in the same nest (polygyny), and colony boundaries appear to be absent with individuals able to move freely between nests (unicoloniality). Such societies depart strongly from a simple family structure and pose a potential challenge to kin selection theory, because high queen number coupled with unrestricted gene flow among nests should result in levels of relatedness among nestmates close to zero. This study investigated the breeding system and genetic structure of a highly polygynous and largely unicolonial population of the wood ant Formica paralugubris. A microsatellite analysis revealed that nestmate workers, reproductive queens and reproductive males (the queens' mates) are all equally related to each other, with relatedness estimates centring around 0.14. This suggests that most of the queens and males reproducing in the study population had mated within or close to their natal nest, and that the queens did not disperse far after mating. We developed a theoretical model to investigate how the breeding system affects the relatedness structure of polygynous colonies. By combining the model and our empirical data, it was estimated that about 99.8% of the reproducing queens and males originated from within the nest, or from a nearby nest. This high rate of local mating and the rarity of long-distance dispersal maintain significant relatedness among nestmates, and contrast with the common view that unicoloniality is coupled with unrestricted gene flow among nests. Received: 8 February 1999 / Received in revised form: 15 June 1999 / Accepted: 19 June 1999