Formica Exsecta Research Articles

Sex Allocation in Mound-Building Ants: The Roles of Resources and Queen Replenishment

Social Hymenoptera have become key organisms for tests of sex-ratio theory. We assess the role of resources for explaining sex-ratio variation in a highly male-biased population of the ant Formica exsecta. Key predictions of two of the three leading hypotheses invoking an effect of resource availability on sex ratios in social insects are not upheld. One prediction of the multifaceted parental-investment hypothesis is that colonies with greater brood production will rear a greater proportion of their diploid offspring as reproductive females (gynes), rather than workers. This study shows the positive correlation between female-biased sex ratio and the production of brood by female-producing colonies was not the outcome of a trade-off in the production of workers vs. gynes. The main prediction of the constant-female hypothesis is that investment in gynes should be constant. We found extreme variation among colonies in the number of gynes produced. By contrast, all the predictions of the queen-replenishment hypothesis were supported. The queen-replenishment hypothesis predicts that colonies produce gynes only when queen number is so low that colony production of brood is reduced, or colony survival threatened. We found that, as predicted by this hypothesis, female-producing colonies had fewer queens and produced a lower biomass of brood than male-producing colonies. In contrast, male-producing colonies had lower production per queen, and were more likely to be limited by external resources. This suggests that queen number limits production in female-producing colonies. These data provide evidence of adaptive adjustment of resources to worker, gyne, and male brood.

Polymorphic microsatellite DNA markers in the ant Formica exsecta

AbstractHighly polymorphic genetic markers provide a useful tool for estimating important genetic parameters in studies of the evolution of sociality in insects. Here we report 14 polymorphic microsatellite markers developed in the ant Formica exsecta. The number of alleles found ranged between 3 and 18 per locus. These markers were developed for studying genetic population structure and mating structure in F. exsecta populations with varying social organizations (monogyne and polygyne types of societies). Cross‐species amplification indicated that some of the markers might be usable even in species belonging to different subfamilies.

Queen recruitment and split sex ratios in polygynous colonies of the ant Formica exsecta

Sex ratios in social insects have become a general model for tests of inclusive fitness theory, sex ratio theory and parent–offspring conflict. In populations of Formica exsecta with multiple queens per colony, sex ratios vary greatly among colonies and the dry‐weight sex ratio is extremely male‐biased, with 89% of the colonies producing males but no gynes (reproductive females). Here we test the queen‐replenishment hypothesis, which was proposed to explain sex ratio specialization in this and other highly polygynous ants (i.e. those with many queens per nest). This hypothesis proposes that, in such ants, colonies produce gynes to recruit them back into the colony when the number of resident queens falls below a given threshold limiting colony productivity or survival. We tested predictions of the queen‐replenishment hypothesis by following F. exsecta colonies across two breeding seasons and relating the change in effective queen number with changes in sex ratio, colony size and brood production. As predicted by the queen‐replenishment hypothesis, we found that colonies that specialized in producing females increased their effective queen number and were significantly more likely to specialize in male production the following year. The switch to male production also coincided with a drop in productivity per queen as predicted. However, adoption of new queens did not result in a significant increase in total colony productivity the following year. We suggest that this is because queen production comes at the expense of worker production and thus queen production leads to resource limitation the following year, buffering the effect of greater queen number on total productivity.

A stratified approach for modeling the distribution of a threatened ant species in the Swiss National Park

We present models predicting the potential distribution of a threatened ant species, Formica exsecta Nyl., in the Swiss National Park (SNP). Data to fit the models have been collected according to a random-stratified design with an equal number of replicates per stratum. The basic aim of such a sampling strategy is to allow the formal testing of biological hypotheses about those factors most likely to account for the distribution of the modeled species. The stratifying factors used in this study were: vegetation, slope angle and slope aspect, the latter two being used as surrogates of solar radiation, considered one of the basic requirements of F. exsecta. Results show that, although the basic stratifying predictors account for more than 50% of the deviance, the incorporation of additional non-spatially explicit predictors into the model, as measured in the field, allows for an increased model performance (up to nearly 75%). However, this was not corroborated by permutation tests. Implementation on a national scale was made for one model only, due to the difficulty of obtaining similar predictors on this scale. The resulting map on the national scale suggests that the species might once have had a broader distribution in Switzerland. Reasons for its particular abundance within the SNP might possibly be related to habitat fragmentation and vegetation transformation outside the SNP boundaries.

Restricted effective queen dispersal at a microgeographic scale in polygynous populations of the ant Formica exsecta.

Ecological constraints on effective dispersal have been suggested to be a key factor influencing social evolution in animal societies as well as the shift from single queen colonies (monogyny) to multiple queen colonies (polygyny) in ants. However, little is known about the effective dispersal patterns of ant queens. Here we investigate the microgeographic genetic structure of mitochondrial haplotypes in polygynous populations of the ant Formica exsecta, both between pastures and among nests within pastures. An analysis of molecular variance revealed a very high genetic differentiation (phiST = 0.72) between pastures, indicating that queens rarely disperse successfully between pastures, despite the fact that pastures were sometimes as close as 1 km. Most of the pastures contained only a single haplotype, and haplotypes were frequently distinct between nearby pastures and even between groups of nests within the same pasture. In the three pastures that contained several haplotypes, haplotypes were not randomly distributed, the genetic differentiation between nests being phiST = 0.17, 0.52, and 0.69. This indicates that most queens are recruited within their parental colonies. However, a large proportion of nests contained more than one haplotype, demonstrating that colonies will sometimes accept foreign queens. The relatedness of mitochondrial genes among nestmates varied between 0.62 and 0.75 when relatedness was measured within each pasture and ranged between 0.72 and 1.0 when relatedness was assessed with all pastures as a reference population. Neighboring nests were more genetically similar than distant ones, and there was significant isolation by distance. This pattern may be due to new nests being formed by budding or by limited effective queen dispersal, probably on foot between neighboring nests. These results show that effective queen dispersal is extremely restricted even at a small geographical scale, a pattern consistent with the idea that ecological constraints are an important selective force leading to the evolution and maintenance of polygyny.

Sex ratio and Wolbachia infection in the ant Formica exsecta.

Sex allocation data in social Hymenoptera provide some of the best tests of kin selection, parent-offspring conflict and sex ratio theories. However, these studies critically depend on controlling for confounding ecological factors and on identifying all parties that potentially manipulate colony sex ratio. It has been suggested that maternally inherited parasites may influence sex allocation in social Hymenoptera. If the parasites can influence sex allocation, infected colonies are predicted to invest more resources in females than non-infected colonies, because the parasites are transmitted through females but not males. Prime candidates for such sex ratio manipulation are Wolbachia, because these cytoplasmically transmitted bacteria have been shown to affect the sex ratio of host arthropods by cytoplasmic incompatibility, parthenogenesis, male-killing and feminization. In this study, we tested whether Wolbachia infection is associated with colony sex ratio in two populations of the ant Formica exsecta that have been the subject of extensive sex ratio studies. In these populations colonies specialize in the production of one sex or the other. We found that almost all F. exsecta colonies in both populations are infected with Wolbachia. However, in neither population did we find a significant association in the predicted direction between the prevalence of Wolbachia and colony sex ratio. In particular, colonies with a higher proportion of infected workers did not produce more females. Hence, we conclude that Wolbachia does not seem to alter the sex ratio of its hosts as a means to increase transmission rate in these two populations of ants.

Colony sex ratios vary with queen number but not relatedness asymmetry in the ant Formica exsecta.

Split-sex-ratio theory assumes that conflict over whether to produce predominately males or female reproductives (gynes) is won by the workers in haplodiploid insect societies and the outcome is determined by colony kin structure. Tests of the theory have the potential to provide support for kin-selection theory and evidence of social conflict. We use natural variation in kinship among polygynous (multiple-queen) colonies of the ant Formica exsecta to study the associations between sex ratios and the relatedness of workers to female versus male brood (relatedness asymmetry). The population showed split sex ratios with about 89% of the colonies producing only males, resulting in an extremely male-biased investment ratio in the population. We make two important points with our data. First, we show that queen number may affect sex ratio independently of relatedness asymmetry. Colonies producing only males had greater genetic effective queen number but did not have greater relatedness asymmetry from the perspective of the adult workers that rear the brood. This lack of a difference in relatedness asymmetry between colonies producing females and those producing only males was associated with a generally low relatedness between workers and brood. Second, studies that suggest support for the relatedness-asymmetry hypothesis based on indirect measures of relatedness asymmetry (e.g. queen number estimated from relatedness data taken from the brood only) should be considered with caution. We propose a new hypothesis that explains split sex ratios in polygynous social insects based on the value of producing replacement queens.

Bears and ants: myrmecophagy by brown bears in central Scandinavia

To determine general patterns of myrmecophagy in bears, we tested hypotheses regarding selection of ant species, factors important to bears when selecting ant species, factors influencing seasonal use of ants, and foraging behavior of brown bears (Ursus arctos) in central Sweden. Ants were an important food for these bears, constituting 12, 16, and 4% of fecal volume in spring, summer, and autumn, respectively. Ants were abundant, 30.5-38.5 tonnes per bear, and bears excavated 8-33% (mean 23%) of the mounds of red forest ants annually. Carpenter ants (Camponotus herculeanus) were highly preferred. Among mound-building red forest ants, the Formica aquilonia/polyctena complex was preferred over Formica exsecta and Formica lugubris. The ants selected by bears had high digestible energy and low formic acid content and behaved passively when the colony was disturbed. Colony size and density may also have influenced the selection of ants. Seasonal use of ants was related not to the availability of pupae or the quality of plant foods but probably to the availability of other foods. Bears consumed only a small proportion of the ants, 4000-5000, each time they opened a mound, probably because of rapidly increasing difficulty in capturing them after the colony was attacked. Eurasian brown bears feed more on ants than North American bears do, perhaps because of greater availability of large colonies of red forest ants. Carpenter ants may have been especially available in our study area following intensive clear-cutting.

Bears and ants: myrmecophagy by brown bears in central Scandinavia

To determine general patterns of myrmecophagy in bears, we tested hypotheses regarding selection of ant species, factors important to bears when selecting ant species, factors influencing seasonal use of ants, and foraging behavior of brown bears (Ursus arctos) in central Sweden. Ants were an important food for these bears, constituting 12, 16, and 4% of fecal volume in spring, summer, and autumn, respectively. Ants were abundant, 30.5-38.5 tonnes per bear, and bears excavated 8-33% (mean 23%) of the mounds of red forest ants annually. Carpenter ants (Camponotus herculeanus) were highly preferred. Among mound-building red forest ants, the Formica aquilonia/polyctena complex was preferred over Formica exsecta and Formica lugubris. The ants selected by bears had high digestible energy and low formic acid content and behaved passively when the colony was disturbed. Colony size and density may also have influenced the selection of ants. Seasonal use of ants was related not to the availability of pupae or the quality of plant foods but probably to the availability of other foods. Bears consumed only a small proportion of the ants, 4000-5000, each time they opened a mound, probably because of rapidly increasing difficulty in capturing them after the colony was attacked. Eurasian brown bears feed more on ants than North American bears do, perhaps because of greater availability of large colonies of red forest ants. Carpenter ants may have been especially available in our study area following intensive clear-cutting.

Sex–ratio regulation: the economics of fratricide in ants

In many insect societies, workers can manipulate the reproductive output of their colony by killing kin of lesser value to them. For instance, workers of the mound-building Formica exsecta eliminate male brood in colonies headed by a single-mated queen. By combining an inclusive fitness model and empirical data, we investigated the selective causes underlying these fratricides. Our model examines until which threshold stage in male brood development do the workers benefit from eliminating males to rear extra females instead. We then determined the minimal developmental stage reached by male larvae before elimination in F. exsecta field colonies. Surprisingly, many male larvae were kept until they were close to pupation, and only then eliminated. According to our model, part of the eliminated males were so large that workers would not benefit from replacing them with new females. Moreover, males were eliminated late in the season, so that new females could no longer be initiated, because matings take place synchronously during a short period. Together, these results indicate that workers did not replace male brood with new females, but rather reduced total brood size during late larval development. Male destruction was probably triggered by resource limitation, and the timing of brood elimination suggests that males may have been fed to females when these start to grow exponentially during the final larval stage. Hence, the evolution of fratricides in ants is best explained by a combination of ecological, demographic and genetic parameters.

Functional groups and patterns of organization in North American ant communities: a comparison with Australia

Ant distribution and behavioural dominance is examined at nine sites along an elevational gradient (1400–2600 m) in south eastern Arizona, in order to classify North American species according to a functional group scheme used extensively in Australia. The functional groups are then used as a basis for determining patterns of community structure along the environmental gradient, and for comparing community structure between Australia and North America. Quantitative information on species com‐ position was obtained from pitfall traps, and patterns of ant abundance at tuna baits were used to determine relative behavioural dominance among taxa. A total of eighty‐three species from twenty‐eight genera was recorded along the elevational gradient, with site species richness ranging from four (high elevation Douglas fir forest) to thirty‐three (mid elevation oak–juniper woodland). There was a strong correlation between ant abundance and richness, which was not an artefact of sampling intensity. The most common ants were species of Forelius, Monomorium, Crematogaster and Pheidole at the three desert sites, species of Formica, Pheidole and Crematogaster at the three woodland sites, and species of Prenolepis and Formica at one forest site. No species were abundant at two other forest sites. The most common species in traps also tended to be the most common species at baits. In terms of behavioural dominance, highly competitive ants included species of Solenopsis, Forelius, Monomorium and Liometopum. Species of Pheidole and Crematogaster tended to be moderately competitive, whereas species of Dory‐ myrmex, Myrmica, Camponotus and Formica (fusca gp) had low competitive ability. On the basis of these results and on published records of other taxa, North American ants were assigned to functional groups as follows (major taxa only given here): Dominant Dolichoderinae—Forelius, Liome‐ topum; Subordinate Camponotini—Camponotus; Hot Climate Specialists—Pogonomyrmex, Myrmecocystus; Cold Climate Specialists—Formica (rufa, exsecta and microgyna groups), Leptothorax, Stenamma, Lasius, Prenolepis; Cryptic Species—Smithistruma, Solenopsis (subgenus Diplorhop‐ trum), Acanthomyops; Opportunists—Formica (fusca group), Myrmica, Paratrechina, Dorymyrmex; Generalized Myr‐ micinae—Pheidole, Crematogaster, Monomorium; Specialist Predators—no major taxa. Functional group composition varied systematically along the elevation gradient: Dominant Dolichoderinae, Generalized MyrÍmicinae and Hot Climate Specialists were predominant at desert sites; Generalized Myrmicinae and Opportunists were predominant at woodland sites; and Opportunists and Cold Climate Specialists were predominant at forest sites. These patterns are consistent with published studies from elsewhere in North America. Almost all North American taxa can be matched with what appear to be ecologically equivalent taxa in Australia, and biogeographic patterns of functional group composition are broadly similar across the two continents. The major differences are that Australian ant communities are far richer in species, and are almost always dominated by dolichoderines, particularly species of Iridomyrmex. Generalized myrmicines are subdominant to dolichoderines in Australia, but are the behaviourally dominant ants throughout the warmer parts of North America. In cool‐temperate North America, species of Formica (especially rufa and exsecta groups) are behaviourally dominant, as they are throughout the Palearctic. Some major features of the North American fauna can be linked to its poor representation of Dominant Dolichoderinae, including (1) the relatively low degree of physiological, morphological and behavioural specialization of Hot Climate Specialists; (2) behavioural dominance by formicines in cool‐temperate habitats; and (3) the susceptibility to invasion by behaviourally dominant species such as the imported fire ant Solenopsis invicta and the Argentine ant Linepithema humile.

Conditional Manipulation of Sex Ratios by Ant Workers: A Test of Kin Selection Theory

Variable queen mating frequencies provide a unique opportunity to study the resolution of worker-queen conflict over sex ratio in social Hymenoptera, because the conflict is maximal in colonies headed by a singly mated queen and is weak or nonexistent in colonies headed by a multiply mated queen. In the wood ant Formica exsecta, workers in colonies with a singly mated queen, but not those in colonies with a multiply mated queen, altered the sex ratio of queen-laid eggs by eliminating males to preferentially raise queens. By this conditional response to queen mating frequency, workers enhance their inclusive fitness.