Population structure and intraspecific aggression in the invasive ant species Anoplolepis gracilipes in Malaysian Borneo

Invasive species are one of the main sources of the ongoing global loss of biodiversity. Invasive ants are known as particularly damaging invaders and their introductions are often accompanied by population-level behavioural and genetic changes that may contribute to their success. Anoplolepis gracilipes is an invasive ant that has just recently received increased attention due to its negative impact on native ecosystems. We examined the behaviour and population structure of A. gracilipes in Sabah, Malaysia. A total of 475 individuals from 24 colonies were genotyped with eight microsatellite markers. Intracolonial relatedness was high, ranging from 0.37 to 1 (mean +/- SD: 0.82 +/- 0.04), while intercolonial relatedness was low (0.0 +/- 0.02, range -0.5-0.76). We compared five distinct sampling regions in Sabah and Brunei. A three-level hierarchical F-analysis revealed high genetic differentiation among colonies within the same region, but low genetic differentiation within colonies or across regions. Overall levels of heterozygosity were unusually high (mean H(O) = 0.95, mean H(E) = 0.71) with two loci being entirely heterozygous, indicating an unusual reproductive system in this species. Bioassays revealed a negative correlation between relatedness and aggression, suggesting kinship as one factor facilitating supercolony formation in this species. Furthermore, we genotyped one individual per nest from Sabah (22 nests), Sarawak (one nest), Brunei (three nests) and the Philippines (two nests) using two mitochondrial DNA markers. We found six haplotypes, two of which included 82.1% of all sequences. Our study shows that the sampled area in Sabah consists of a mosaic of differently interrelated nests in different stages of colony establishment. While some of the sampled colonies may belong to large supercolonies, others are more likely to represent recently introduced or dispersed propagules that are just beginning to expand.     

Mating frequency and genetic structure of the Argentine ant Linepithema humile

The nest and population genetic structures of the Argentine ant, Linepithema humile were investigated using eight microsatellite loci. Genotypes of the sperm from spermathecae of 87 queens were consistent with all queens being singly inseminated. The probability of a double mating remaining undetected was low (0.012) suggesting that no queens or only a very low proportion mate multiply. The relatedness between the queens and their mates was negative (R = -0.164 +/- 0.044) and significantly different to zero (P = 0.020). However, the high negative relatedness value was caused by a significant allele frequency difference between the sexes at a single locus (Lhum-28). When this locus was removed from the analyses, the relatedness was not significantly different from zero (R = 0.013 +/- 0.050, P = 0.812). Analysis of 10 nests revealed that the genetic differentiation among nests was weak (FST = 0.003) and not distinguishable from zero (P = 0.468). Similarly, the overall relatedness among nestmate females was not significantly different from zero (R = 0.007 +/- 0.018, P = 0.706). These results are consistent with the lack of distinct nest boundaries and the large number of queens per nest in the population studied. Although mating takes place inside the nest, the inbreeding coefficient was close to zero (F = 0.007 +/- 0.025, P = 0.786). Overall, these data indicate substantial local gene flow mediated by movement of reproductives among colonies.     

Why Wasp Foundresses Change Nests: Relatedness,Dominance, and Nest Quality

The costs and benefits of different social options are best understood when individuals can be followed as they make different choices, something that can be difficult in social insects. In this detailed study, we follow overwintered females of the social wasp Polistes carolina through different nesting strategies in a stratified habitat where nest site quality varies with proximity to a foraging area, and genetic relatedness among females is known. Females may initiate nests, join nests temporarily or permanently, or abandon nests. Females can become helpers or egglayers, effectively workers or queens. What they actually do can be predicted by a combination of ecological and relatedness factors. Advantages through increased lifetime success of individuals and nests drives foundresses of the social wasp Polistes from solitary to social nest founding. We studied reproductive options of spring foundresses of P. carolina by monitoring individually-marked wasps and assessing reproductive success of each foundress by using DNA microsatellites. We examined what behavioral decisions foundresses make after relaxing a strong ecological constraint, shortage of nesting sites. We also look at the reproductive consequences of different behaviors. As in other Polistes, the most successful strategy for a foundress was to initiate a nest as early as possible and then accept others as subordinates. A common feature for many P. carolina foundresses was, however, that they reassessed their reproductive options by actively monitoring other nests at the field site and sometimes moving permanently to new nests should that offer better (inclusive) fitness prospects compared to their original nests. A clear motivation for moving to new nests was high genetic relatedness; by the end of the foundress period all females were on nests with full sisters.     

Colony structure of a slavemaking ant. I. Intracolony relatedness, worker reproduction, and polydomy

Colony and population structure of the obligate slavemaker ant Protomognathus americanus was analyzed via four nuclear microsatellite loci and mitochondrial DNA (mtDNA) markers. Colonies of P. americanus usually contain a single queen, and here we show that she is singly inseminated. Nestmate workers are generally full sisters and their relatedness does not deviate from the expected value of 0.75. Even though colonies were strictly monogynous, we were able to infer that colony takeover by related queens was common and queen replacement by unrelated queens was rare. Polydomy is widespread, with neighboring nests having the same genetic composition. Although we found no evidence of population viscosity or inbreeding from nuclear markers, mtDNA markers provided evidence for small-scale genetic structuring. Haplotype structuring and takeover by related queens suggest philopatry of newly mated queens. In this species, workers reproduce in queenright and queenless nests and worker reproduction accounts for more than 70% of all males. Although sex-ratio theory points to slavemaking ants as important systems for studying queen-worker conflict, our results indicate no basis for such conflict in P. americanus, because extensive worker reproduction generates shifts in relatedness values. Rather, the dual effects of independent polydomous nest units and local resource competition among queens produce male-biased allocation ratios in this species.     

Plasticity in queen number and social structure in the invasive Argentine ant (Linepithema humile)

In many polygynous social insect societies, ecological factors such as habitat saturation promote high queen numbers by increasing the cost of solitary breeding. If polygyny is associated with constrained environments, queen number in colonies of invasive social insects should increase as saturation of their new habitat increases. Here I describe the variation in queen number, nestmate relatedness, and nest size along a gradient of time since colonization in an invading population of Argentine ants (Linepithema humile) in Haleakala, Hawaii. Nest densities in this population increase with distance from the leading edge of the invasion, reaching a stable density plateau approximately 80 m from the edge (> 2 years after colonization). Although the number of queens per nest in Haleakala is generally lower than previously reported for Argentine ants, there is significant variation in queen number across this population. Both the observed and effective queen numbers increase across the density gradient, and nests in the center of the population contain queen numbers three to nine times higher than those on the edge of the invasion. The number of workers per nest is correlated with queen number, and nests in the center are six times larger than nests at the edge. Microsatellite analysis of relatedness among nestmates reveals that all nests in the Haleakala population are characterized by low relatedness and have evidence of multiple reproducing queens. Relatedness values are significantly lower in nests in the center of the population, indicating that the number of reproducing queens is greater in areas of high nest density. The variation in queen number and nestmate relatedness in this study is consistent with expectations based on changes in ecological constraints during the invasion of a new habitat, suggesting that the social structure of Argentine ant populations is strongly influenced by ecological factors. Flexibility in social structure may facilitate persistence in variable environments and may also confer significant advantages to a species when introduced into new areas.     

Colony structure of a slavemaking ant. II. Frequency of slave raids and impact on the host population

The parasite pressure exerted by the slavemaker ant Protomognathus americanus on its host species Leptothorax longispinosus was analyzed demographically and genetically. The origin of slaves found in colonies of the obligate slavemaker was examined with nuclear and mitochondrial DNA markers to make inferences about the frequency and severity of slave raids. Relatedness of enslaved L. longispinosus workers in the same nest was very low, and our data suggest that, on average, each slavemaker nest raids six host colonies per season. Therefore, the influence of slavemaker species on their hosts is much stronger than simple numerical ratios suggest. We also found that slave relatedness was higher in small than in large slavemaker nests; thus, larger nests wield a much stronger influence on the host. We estimated that in the study population, on average, a host nest has a 50% chance of being attacked by a slavemaker colony per year. Free-living Leptothorax colonies in the vicinity of slavemaker nests did not represent the source of slaves working in P. americanus colonies, which suggests that raided nests either do not survive or migrate after being raided. Colony composition and intranest relatedness of free-living L. longispinosus colonies differed markedly between areas with slavemakers and those that are parasite-free. In the presence of slavemakers, host colonies were less likely to be polygynous and had fewer workers and a higher relatedness among worker brood. Host nests with slavemaker neighbors allocated more resources into sexuals, possibly caused by these shifts in nest demography. Finally, enslaved Leptothorax workers in P. americanus nests appeared to be less efficient than their counterparts in free-living colonies. Thus, slavemakers exert a much stronger impact on their hosts than had previously been suspected and represent an unique system to study parasite-host coevolution.     

Host-parasite kinship in a female-philopatric bird population: evidence from relatedness trend analysis

Conspecific brood parasitism (CBP), an alternative reproductive tactic where some females lay eggs in the nests of other females of the same species, occurs in many animals with egg care. It is particularly common in waterfowl, for reasons that are debated. Many waterfowl females nest near their birthplace, making it likely that some local females are relatives. We analyse brood parasitism in a Hudson Bay population of common eiders, testing predictions from two alternative hypotheses on the role of relatedness in CBP. Some models predict host-parasite relatedness, others predict that parasites avoid close relatives as hosts. To distinguish between the alternatives, we use a novel approach, where the relatedness of host-parasite pairs is tested against the spatial population trend in pairwise relatedness. We estimate parasitism, nest take-over and relatedness with protein fingerprinting and bandsharing analysis of egg albumen, nondestructively sampled from each new egg in the nest throughout the laying period. The results refute the hypothesis that parasites avoid laying eggs in the nests of related hosts, and corroborate the alternative of host-parasite relatedness. With an estimated r of 0.12-0.14, females laying eggs in the same nest are on average closer kin than nesting neighbour females. Absence of a population trend in female pairwise relatedness vs. distance implies that host-parasite relatedness is not only an effect of strong natal philopatry: some additional form of kin bias is also involved.     

Living with strangers: direct benefits favour non-kin cooperation in a communally nesting bird

The greater ani (Crotophaga major), a Neotropical cuckoo, exhibits an unusual breeding system in which several socially monogamous pairs lay eggs in a single nest and contribute care to the communal clutch. Cooperative nesting is costly-females compete for reproduction by ejecting each other's eggs-but the potential direct or indirect fitness benefits that might accrue to group members have not been identified. In this study, I used molecular genotyping to quantify patterns of genetic relatedness and individual reproductive success within social groups in a single colour-banded population. Microsatellite analysis of 122 individuals in 49 groups revealed that group members are not genetic relatives. Group size was strongly correlated with individual reproductive success: solitary pairs were extremely rare and never successful, and nests attended by two pairs were significantly more likely to be depredated than were nests attended by three pairs. Egg loss, a consequence of reproductive competition, was greater in large groups and disproportionately affected females that initiated laying. However, early-laying females compensated for egg losses by laying larger clutches, and female group members switched positions in the laying order across nesting attempts. The greater ani, therefore, appears to be one of the few species in which cooperative breeding among unrelated individuals is favoured by direct, shared benefits that outweigh the substantial costs of reproductive competition.     

Egg morphology fails to identify nests parasitized by conspecifics in common pochard: a test based on protein fingerprinting and including female relatedness

Conspecific brood parasites lay eggs in nests of other females of the same species. A variety of methods have been developed and used to detect conspecific brood parasitism (CBP). Traditional methods may be inaccurate in detecting CBP and in revealing its true frequency. On the other hand more accurate molecular methods are expensive and time consuming. Eadie developed a method for revealing CBP based on differences in egg morphology. That method is based on Euclidean distances calculated for pairs of eggs within a clutch using standardized egg measurements (length, width and weight). We tested the applicability of this method in the common pochard Aythya ferina using nests that were identified as parasitized (39 nests) or non‐parasitized (16 nests) based on protein fingerprinting of eggs. We also analyzed whether we can distinguish between parasitic and host eggs in the nest. We found that variation in MED can be explained by parasitism but there was a huge overlap in MED between parasitized and non‐parasitized nests. MED also increased with clutch size. Using discriminant function analysis (DFA) we found that only 76.4% of nests were correctly assigned as parasitized or non‐parasitized and only 68.3% of eggs as parasitic or host eggs. Moreover we found that MED in parasitized nests increased with relatedness of the females that laid eggs in the nest. This finding was supported by positive correlation between MED and estimated relatedness in female–female pairs. Although variation in egg morphology is associated with CBP, it does not provide a reliable clue for distinguishing parasitized nests from non‐parasitized nests in common pochard.     

Relatedness and genetic structure in a socially polymorphic population of the spider Anelosimus studiosus

The evolution of sociality remains a challenge in evolutionary biology and a central question is whether association between kin is a critical factor favouring the evolution of cooperation. This study examines genetic structure of Anelosimus studiosus, a spider exhibiting polymorphic social behaviour. Two phenotypes have been identified: an ‘asocial’ phenotype with solitary female nests and a ‘social’ phenotype with multi‐female/communal nests. To address the questions of whether these phenotypes are differentiated populations and whether cooperative individuals are closely related, we used microsatellites to analyse individuals from both communal and solitary nests. We found no evidence of differentiation between social and solitary samples, implying high rates of interbreeding. This is consistent with the hypothesis that these phenotypes coexist as a behavioural polymorphism within populations. Pairwise relatedness coefficients were used to test whether cooperating individuals are more closely related than expected by chance. Pairwise relatedness of females sharing communal webs averaged 0.25, the level expected for half‐siblings and significantly more closely related than random pairs from the population. Solitary females collected at similar distances to the communal spider pairs were also more closely related than expected by chance (mean relatedness = 0.18), but less related than social pairs. These results imply that low dispersal contributes to increase likelihood of interaction between kin, but relatedness between social pairs is not explained by spatial structure alone. We propose that these phenotypes represent stages in the evolution of sociality, where viscous population structure creates opportunities for kin selection and cooperation is favoured under certain environmental conditions.     

RELATEDNESS AND POPULATION STRUCTURE OF THE PRIMITIVELY EUSOCIAL BEE LASIOGLOSSUM ZEPHYRUM (HYMENOPTERA: HALICTIDAE) IN KANSAS

Lasioglossum zephyrum is a primitively eusocial bee, which nests in small colonies of up to 20 individuals. The nests occur in patchily distributed aggregations of from a few to over 1,000 nests along periodically disturbed stream and river banks in eastern North America. We used five polymorphic allozyme loci to test for geographic structure and estimate relatedness in eight patches of nests from five aggregations in Douglas Co., Kansas. Autocorrelation analysis of gene frequencies, plus a multilocus G test, revealed a low but significant tendency for differentiation among nests within patches, among patches within aggregations, and among aggregations. Small numbers of nests restricted estimation of relatedness to three patches, of which only one had a sample size large enough to yield confidence limits narrow enough to be informative. The limits from this patch of 20 nests are 0.64 < 0.8245 < 1.01. While these limits are consistent with the true value being 0.75 (that expected under male-haploidy if each nest results from the reproduction of a single, once-mated female), the occurrence of some nests with three or more genotypes shows that nest makeup is more complex than this, so that a lower value, say 0.7, is more plausible. This value is sufficiently high to indicate that kin selection is probably important in these populations.     

Clumping versus spacing out: Experiments on nest predation in fieldfares (Turdus pilaris)

Fieldfares (Turdus pilaris), which nest solitarily as well as in colonies, offer an opportunity to assess within one species the relative advantages of clumping versus spacing out. An experiment with artificial, egg-baited nests showed that predation in the absence of fieldfares was higher on clumped than on scattered nests. In another experiment, we tested whether artificial nests run a higher risk of predation near solitary fieldfare pairs than near fieldfare colonies. Nest predation was higher near solitary fieldfare pairs than outside and inside fieldfare colonies. The risk probably was higher outside than inside colonies. Communal antipredator attacks is the likely cause of reduced predation near colonies. Even solitary fieldfare pairs confer some degree of protection, because predation was lower at artificial nests near solitary fieldfare pairs than at nests without neighbouring fieldfares.     

PVC nest boxes are less at risk of occupancy by feral honey bees than timber nest boxes and natural hollows

Feral European Honey Bee (Apis mellifera) has been identified as a potential nest competitor for Australian hollow nesting species, but few studies have investigated the impact of feral honey bee competition on Threatened species. Our study used data from Glossy Black‐cockatoo (Calyptorhynchus lathami halmaturinus) nests on Kangaroo Island, monitored and managed over an 11‐year period, and found 12% of nests became occupied by feral honey bees during that period. Our results indicate that feral honey bees were less likely to occupy nest boxes made of PVC (5%) compared with wooden nest boxes (24%) or natural hollows in Eucalyptus trees (14%). The removal of feral honey bee hives from nests is a priority for long‐term conservation of glossy black‐cockatoos on Kangaroo Island. We recommend that PVC nest boxes are chosen for future nesting habitat restoration, due to the more frequent use of wooden nest boxes by feral honey bees.     

Breeding behaviour, relatedness and sex-investment ratios inLeptothorax tuberum Fabricius

The ant speciesLeptothorax tuberum was shown to be predominantly monogynous. Queens usually mate once only but some nests may have a multiply-mated queen or are partially or serially polygynous. As expected from these results, within nest relatedness between workers and between workers and alate queens was found to be high. Almost fifty percent of nests had no nest queen which may indicate high queen mortality, queens leaving to found new nests or nest fragmentation. Observed female investment frequencies (IF _o ) were not significantly different from those expected on the basis of worker control of sexual production and the relatedness estimates of workers to alate queens and workers to males calculated from isozyme data (IF _E ). These values were not consistent with queen control. There was no evidence for lower IF _o s in queenless nests nor for higher IF _o s in larger nests classified by worker number. When nests were classified by sexual productivity, however, there was a strong rank correlation between productivity and female bias. This is the first study of an ant species to test observed IF against expected IF calculated without inferring between caste relatedness from worker data or pooling of data from different castes.     

Shared reproduction and sex ratio adjustment to clutch size in a socially polymorphic orchid bee

Nests of the primitively eusocial orchid bee Euglossa viridissima are generally founded by a solitary female but can be reactivated by female offspring, in the presence or absence of the mother. The population therefore exists of solitary and social nests that co-occur in an area. A female can stay as a subordinate helper under a dominant female or disperse to become a solitary foundress. Yet, the costs and benefits of the different social phenotypes are so far little understood. Here, we compared solitary and social nests of orchid bees. We used offspring of solitary and social nests to calculate offspring sex ratio, and applied molecular markers to calculate intranidal relatedness, infer maternity and test whether sociality may have a genetic component. We found that social nests had on average more brood than solitary nests. The overall sex ratio in the population did not differ from 1:1. However, social nests tended to produce a split sex ratio with some nests producing mainly males and others mainly female offspring. Regardless of social phenotype, the number of offspring was correlated with the sex ratio, with smaller nests having a more female-biased offspring. In social nests, not all offspring resulted from a single-mated mother, which was also the case for some solitary nests. This suggests shared reproduction in social nests and may be an indication that intraspecific parasitism and nest takeover are not uncommon. Structure analyses did not reveal different genetic background of the two social phenotypes. Our results suggest that there is no clear benefit that favours one of the two social phenotypes over the other and that the population is kept at balance in terms of sex ratio.     

Repeatability of nest predation in passerines depends on predator species and time scale

It has been proposed that some specific locations of bird's nests have higher intrinsic chances of being depredated than other locations. This predicts that fates of consecutive nesting attempts at the same site should be repeatable. We used 20 pairs of old thrush nests to simulate repeated nesting attempts at the same sites, both within and between breeding seasons (n=40  sites×2  trials×2  years=160). Each nest was monitored by a camera to record multiple predation events and to identify predators. Predation by all predator species was repeatable during a 15-day trial. Predation by principal predators (jay Garrulus glandarius , marten Martes martes / foina ) and total predation (all species combined) was not correlated within pairs of simultaneously exposed nests or within samples of nests from particular study plot, and not repeatable for individual nests between-trials or between-years. These findings suggest short-term effect of predator memory causing revisitation of previously depredated nests during a current nesting trial (all predators); do not support an effect of nest site features on multiple nest discoveries and/or an effect of nest location on repeated random encounters with the same nest (principal predators). Long-term repeatability and correlation within pairs of simultaneously exposed nests was detectable only in occasional predators (great spotted woodpecker Dendrocopos major , possibly also squirrel Sciurus vulgaris ), which suggests effect of nest location combined with site fidelity and individual foraging specialization of these predators. We conclude that repeatability of nest predation depends on the time scale considered and the local predator community. We caution against spurious findings of repeatable nest predation resulting simply from statistical properties of correlation in binary data (nest fates).     

Nest Suitability,Fine-Scale Population Structure and Male-Mediated Dispersal of a Solitary Ground Nesting Bee in an Urban Landscape

Bees are the primary pollinators of flowering plants in almost all ecosystems. Worldwide declines in bee populations have raised awareness about the importance of their ecological role in maintaining ecosystem functioning. The naturally strong philopatric behavior that some bee species show can be detrimental to population viability through increased probability of inbreeding. Furthermore, bee populations found in human-altered landscapes, such as urban areas, can experience lower levels of gene flow and effective population sizes, increasing potential for inbreeding depression in wild bee populations. In this study, we investigated the fine-scale population structure of the solitary bee Colletes inaequalis in an urbanized landscape. First, we developed a predictive spatial model to detect suitable nesting habitat for this ground nesting bee and to inform our field search for nests. We genotyped 18 microsatellites in 548 female individuals collected from nest aggregations throughout the study area. Genetic relatedness estimates revealed that genetic similarity among individuals was slightly greater within nest aggregations than among randomly chosen individuals. However, genetic structure among nest aggregations was low (Nei’s G_ST = 0.011). Reconstruction of parental genotypes revealed greater genetic relatedness among females than among males within nest aggregations, suggesting male-mediated dispersal as a potentially important mechanism of population connectivity and inbreeding avoidance. Size of nesting patch was positively correlated with effective population size, but not with other estimators of genetic diversity. We detected a positive trend between geographic distance and genetic differentiation between nest aggregations. Our landscape genetic models suggest that increased urbanization is likely associated with higher levels of inbreeding. Overall, these findings emphasize the importance of density and distribution of suitable nesting patches for enhancing bee population abundance and connectivity in human dominated habitats and highlights the critical contribution of landscape genetic studies for enhanced conservation and management of native pollinators.     

Inquilines of Brachymenes dyscherus nests with special reference to Monobia schrottkyi (Hymenoptera,Vespidae, Sphecidae)

Sixty-four inactive nests of the solitary mud-daubing wasp Brachymenes dyscherus, reused by 5 inquiline species, were collected at Fazenda Santa Carlota, Cajuru, S?o Paulo, Brazil in 1995 and 1996. Monobia schrottkyi used 52 nests; among the 717 cells available for use, 502 were reused. The number of cells per nest varied from 3 to 24; 1 to 16 individuals emerged from September to April (154 males and 112 females). Forty-six cells were parasitized by Melittobia sp. (n = 44) and Ichneumonidae (n = 2). Monobia curvata used 3 nests; among the 50 cells available for use, 38 were reused and 15 males and 8 females emerged from August to November. Three cells were parasitized by Ichneumonidae. Montezumia petiolata occupied 1 nest; among the 8 available cells, 7 were reused and 2 males and 3 females emerged in September. Podium denticulatum used 2 nests; the 20 cells available for use were reused and 11 males and 4 females emerged in August. Trypoxylon rogenhoferi used 5 nests that had 65 available cells; 48 of them were reused, from which 19 males and 11 females emerged from September to November. Three cells were parasitized by Ichneumonidae (n = 2) and Chrysididae (n = 1).     

Queen recruitment in a multiple-queen population of the fire ant Solenopsis invicta

We assessed patterns of new queen recruitment in a polygyne(multiple queens per nest) population of the fire ant Solenopsisinvicta in its introduced range. Newly recruited queens wereidentified using four physiological markers, and genotypic datafrom nuclear and mitochondrial markers were used to estimaterelatedness of new nest mate queens to each other and to theolder nest mate queens. In total, 1.7% of the queens collectedin late spring and early summer were deemed to be new recruits.The relatedness of these queens to each other and to the olderqueens within nests was not significantly different from zero,suggesting that newly recruited queens represent a random sampleof potential reproductive queens in the population. Moreover,the number of new queens recruited within nests was not correlatedwith the number of older queens present and did not differ significantlyfrom a Poisson distribution. Thus, queen recruitment in this populationof S. invicta appears to occur at random with respect to thenumber of older queens present within nests.     

Highly variable social organisation of colonies in the ant Formica cinerea

Social organisation of colonies was examined in the ant Formica cinerea by estimating the coefficient of genetic relatedness among worker nest mates. The estimates based on microsatellite genotypes at three loci ranged from values close to zero to 0.61 across the populations studied in Finland. These results showed that a fundamental feature of colonies, the number of reproductive queens, varied greatly among the populations. Colonies in some populations had a single queen, whereas the nests could have a high number number of queens in other populations. There was a weak but non-significant correlation between the genetic and metric distance of nests within two populations with intermediate level of relatedness. Differentiation among nearby populations (within the dispersal distance of individuals) in one locality indicated limited dispersal or founder effects. This could occur when females are philopatric and stay in the natal polygynous colony which expands by building a network of nest galleries within a single habitat patch.