Intranest Relatedness Research Articles

Colony structure, population structure, and sharing of foraging trees in the ant Myrmecia nigriceps (Hymenoptera: Formicidae)

Foraging ants face many dangers in search of food and often need to defend their prey to ensure the colony’s survival, although ants may also follow a peaceful foraging strategy. A non-aggressive approach is seen in the Australian bull ant Myrmecia nigriceps, in that workers of neighboring nests sometimes share foraging trees. In this study, we observed 31 nests at Mount Majura Nature Reserve in Canberra (Australia), 12 of which shared a foraging tree with at least one other nest in at least one of three nights. We genotyped 360 individuals at five published microsatellite loci and further established a set of nine polymorphic loci for M. nigriceps. Our results revealed a significant correlation between tree sharing and geographical distance between nests. We found no correlation between internest relatedness and tree sharing, geographical distance between nests and internest relatedness, and intranest relatedness and tree sharing. We further investigated the colony structure of M. nigriceps. All colonies were monodomous; the number of queens per colony ranged from one to two, and the number of fathers from one to three. No instances of worker drifting were found in this study.

Kinship, parental manipulation and evolutionary origins of eusociality.

One of the hallmarks of eusociality is that workers forego their own reproduction to assist their mother in raising siblings. This seemingly altruistic behaviour may benefit workers if gains in indirect fitness from rearing siblings outweigh the loss of direct fitness. If worker presence is advantageous to mothers, however, eusociality may evolve without net benefits to workers. Indirect fitness benefits are often cited as evidence for the importance of inclusive fitness in eusociality, but have rarely been measured in natural populations. We compared inclusive fitness of alternative social strategies in the tropical sweat bee, Megalopta genalis, for which eusociality is optional. Our results show that workers have significantly lower inclusive fitness than females that found their own nests. In mathematical simulations based on M. genalis field data, eusociality cannot evolve with reduced intra-nest relatedness. The simulated distribution of alternative social strategies matched observed distributions of M. genalis social strategies when helping behaviour was simulated as the result of maternal manipulation, but not as worker altruism. Thus, eusociality in M. genalis is best explained through kin selection, but the underlying mechanism is likely maternal manipulation.

Locally adapted social parasite affects density, social structure, and life history of its ant hosts

Selection and adaptation are important processes in the coevolution between parasites and their hosts. The slave-making ant Protomognathus americanus, an obligate ant social parasite, has previously been shown to evolve morphological, behavioral, and chemical adaptations in the coevolutionary arms race with its Temnothorax hosts. Yet empirical studies have given variable results on the strength of the selection pressure this parasite exerts on its host populations. In this study, we directly investigated the pressure exerted by P. americanus and the reactions of the main host species, T. longispinosus, in two ant communities by manipulating parasite density in the field over several years. In addition, a cross-fostering design with the exchange of parasites between host populations allowed us to investigate local adaptation of parasite or host. We demonstrate a severe impact of the social parasite on the two host populations in West Virginia and New York, but also variation in host reactions between sites, as expected by the geographic mosaic theory of coevolution. Host density decreased at the West Virginia site with the presence of local slave-makers, whereas at the ecologically favorable New York site, density was unaffected. Nevertheless, social organization, colony size, and investment patterns of these host colonies at this site changed in response to our parasite manipulation. The release of P. americanus colonies led to a reduction in the number of resident queens and workers, an increase in intranest relatedness, and lower productivity, but also a higher investment in reproductives. In West Virginia, colony demography did not change, but raiding activity by New York slave-makers caused different investment patterns of host colonies. In addition, the cross-fostering element revealed local adaptation of the parasite P. americanus: slave-making colonies fared better in their sympatric host population, as they contained more slave-making ant workers and slaves at the end of our 27-month experiment.

Kin recognition and the paradoxical patterns of aggression between colonies of a Mojave desert Pheidole ant

Populations of the desert seed-harvesting ant Phei- dole xerophylla are often characterized by high nest density leading to competitive interactions between foragers from different nests. We investigated the inter-nest aggression, spatial distribution and genetic structure of a P. xerophylla population of the Mojave Desert in Southern California. Inter-nest aggression was quantifi ed by standardized staged encounters in a neutral arena. Genetic relatedness within nests and relatedness between nests were calculated using al- lelic frequencies at four microsatellite-DNA loci. We found a bimodal distribution of inter-colony aggression levels with a fi rst mode at low aggression levels and another mode at much higher aggression levels. Inter-colony aggression levels were largely non-transitive. No effect of geographical distance on inter-nest aggression levels was detected. De- spite high amounts of variation in inter-colony relatedness (-0.24 to 0.37) this variable did not correlate with the level of aggression between nests. Intra-nest relatedness ranged from 0.40 to 0.75 and close inspection of worker genotypes within colonies revealed a high proportion of polygynous colonies or a mixture of polygyny and polyandry. Aggression levels among nests was found to decrease with increasing intra-nest relatedness. These results do not support the idea that ag- gression is modulated by a nestmate recognition mechanism based on overall genetic similarity. Instead, the absence of transitivity found in inter-colony aggression and bimodal distribution of aggression levels are compatible with a com- mon label acceptance model of nestmate recognition and suggest that label diversity may be encoded by a limited number of loci.

Seasonal polydomy and unicoloniality in a polygynous population of the red wood ant Formica truncorum

Ant colonies may have a single or several reproductive queens (monogyny and polygyny, respectively). In polygynous colonies, colony reproduction may occur by budding, forming multinest, polydomous colonies. In most cases, budding leads to strong genetic structuring within populations, and positive relatedness among nestmates. However, in a few cases, polydomous populations may be unicolonial, with no structuring and intra-nest relatedness approaching zero. We investigated the spatial organisation and genetic structure of a polygynous, polydomous population of Formica truncorum in Finland. F. truncorum shifts nest sites between hibernation and the reproductive season, which raises the following question: are colonies maintained as genetic entities throughout the seasons, or is the population unicolonial throughout the season? Using nest-specific marking and five microsatellite loci, we found a high degree of mixing between individuals of the population, and no evidence for a biologically significant genetic structuring. The nestmate relatedness was also indistinguishable from zero. Taken together, the results show that the population is unicolonial. In addition, we found that the population has undergone a recent bottleneck, suggesting that the entire population may have been founded by a very limited number of females. The precise causes for unicoloniality in this species remain open, but we discuss the potential influence of intra-specific competition, disintegration of recognition cues and the particular hibernation habits of this species.

Ecology of Leptothorax 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.

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.

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.

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.

Migration between nests in the Australian arid‐zone ant Rhytidoponera sp. 12 revealed by DGGE analyses of mitochondrial DNA

Denaturing gradient gel electrophoresis (DGGE) of a PCR‐amplified region of the mitochondrial DNA (mtDNA) including a large part of the cytochrome b gene revealed four haplotypes among worker ants from 75 colonies of the queenless ant Rhytidoponera sp. 12. The DGGE results were checked by sequencing examples of the haplotypes; all changes were transitions and two haplotypes differed by only a single substitution. Previous work (e.g. Crozier et al. 1984) showed that intranest relatedness is low for nuclear genes yet neighbouring nests are related; gene flow via winged males appeared the best explanation for this phenomenon. Two mtDNA haplotypes were found in 34.7% of the colonies studied, showing that female movement also occurs between nests. Migration of mated individuals on such a large scale when the number of reproductives is relatively small is unexpected (Crozier & Pamilo 1996). An observed tendency to clumping of the haplotypes is in accordance with the wingless nature of the female dispersers.

Nestmate discrimination in a ponerine ant (Rhytidoponera sp. 12) without a queen caste and with a low intra-nest relatedness

Various ponerine ants do not have a queen caste, and it is not known whether this modification in the eusocial structure affects the ability to recognize between nestmates and non-nestmates. Foragers ofRhytidoponera sp. 12 were collected from 15 adjacent nests, and were immediately tested in pairwise introductions between nests. Out of 179 tests, 84 % resulted in fighting, and in another 5 % there were unambiguous signs of hostility. Furthermore, natural aggression between marked foreign workers was observed near the nests. There are many mated laying workers in the colonies ofR. sp. 12, and this leads to nestmates being less genetically-similar (whereupon their genotypically-correlated recognition odours may vary considerably). Since there is no influence of a queen odour, the existence of a distinct colony identity reveals that workers are able to learn the spectrum of individual odours of their nestmates. In instances where workers from two foreign nests did not fight, each of them was again tested with other foreign ants (from the same or a different nest). They could discriminate in some cases but not others. This suggests that a gestalt odour does not exist in these colonies, but larger samples are needed to confirm this. Colonies ofR. sp. 12 are not polydomous, as had been previously suggested. The transfer of workers which sometimes occurs between nests represents episodes of nest emigration.

Genetic organization of Formica sanguinea populations

1. Genetic organization of two populations (Bembole and Skomakarskar) of the ant Formica sanguinea was studied using a gene locus encoding malate dehydrogenase enzyme as a marker gene. Five alleles were found segregating at the locus. 2. There was a slight excess of heterozygous gynes in the Bembole population, but the pooled genotype frequencies in the workers were close to the expected Hardy-Weinberg ratios. Genetic relatedness among single-nest gynes (0.21) was of the same magnitude as that among worker nest mates (0.19). This suggests that the cooperating gynes may have arisen from a single nest. On the basis of the relatedness analysis, the mean estimate for the gyne number per nest was 5.3 in the Bembole and 1.9 in the Skomakarskar population. 3. There was extensive microdifferentiation within the Skomakarskar population, the frequencies of the alleles b and c changing greatly within a distance of some tens of meters (Fig. 1). 4. Genetic organizations of the Bembole and Skomakarskar populations differed from each other with respect to intranest heterogeneity and affinity of nearest neighbor nests (Fig.2). Genetic heterogeneity within single nests was greater at Bembole, as revealed both by lower relatedness of nest mates and by a greater number of multigenotype nests. The neighboring nests showed clear positive relatedness to each other on Skomakarskar (0.24) but not at Bembole (-0.06). The results fit the assumption that the threshold for nest splitting is higher at Bembole, leading to a higher degree of polygyny, a lower level of intranest relatedness, and to a weak tendency of daughternest formation.