Foraging and spatiotemporal territories in the honey ant Myrmecocystus mimicus wheeler (Hymenoptera: Formicidae)

Summary The honey ant Myrmecocystus mimicus is a scavenger, forages extensively on termites, collects floral nectar, and tends homoptera. Individual foragers of M. mimicus usually disperse in all directions when leaving the nest, but there are also groups of foragers that tend to swarm out of the nest primarily in one direction. Such massive departues are usually at irregular intervals, which may last several hours. The results of field and laboratory experiments suggest that these swarms of foragers are organized by a group recruitment process, during which recruiting scout ants lay chemical orientation trails with hindgut contents and simultaneously stimulate nestmates with a motor display and secretions from the poison gland. Usually these columns travel considerable distances (4–48 m) away from the nest, frequently interfering with the foraging activity of conspecific neighboring colonies.To prevent a neighboring colony from access to temporal food sources or to defend spatiotemporal borders, opposing colonies engage in elaborate display tournaments. Although hundreds of ants are often involved during these tournaments almost no physical fights occur. Instead, individual ants confront each other in highly sterotyped aggressive displays, during which they walk on stilt legs while raising the gaster and head. Some of the ants even seem to inflate their gasters so that the tergites are raised and the whole gaster appears to be larger. In addition, ants involved in tournament activities are on average larger than foragers.The dynamics of the tournament interactions were observed in several colonies over several weeks-mapping each day the locations of the tournaments, the major directions of worker routes away from the nest, and recording the general foraging activities of the colonies. The results indicate that a kind of dominance order can occur among neighboring colonies. On the other hand, often no aggressive interactions among neighboring colonies can be observed, even though the colonies are actively foraging. In those cases the masses of foragers of each colony depart in one major direction that does not bring them into conflict with the masses of foragers of a neighboring colony. This stability, however, can be disturbed by offering a new rich food source to be exploited by two neighboring colonies. This invariably leads to tournament interactions.When a colony is considerably stronger than the other, i.e., with a much larger worker force, the tournaments end quickly and the weaker colony is raided. The foreign workers invade the nest, the queen of the resident colony is killed or dirven off, while the larvae, pupae, callow workers, and honey pot workers are carried or dragged to the nest of the raiders. From these and other observations we conclude that young M. mimicus queens are unlikely to succeed in founding a colony within approximately 3 m of a mature M. mimicus colony because they are discovered and killed, or driven off by workers of the resident colony. Within approximately 3–15 m queens are more likely to start colonies, but these incipient groups run a high risk of being raided and exterminated by the mature colony.Although populations of M. mimicus and M. depilis tend to replace each other, there are areas where both species overlap marginally. Foraging areas and foraging habitats of both species also overlap broadly, but we never observed tournament interactions between M. mimicus and M. depilis.The adaptive significance of the spatiotemporal territories in M. mimicus is discussed.     

Polygyny,relatedness and nest founding in the polygynous myrmicine ant Leptothorax acervorum (Hymenoptera; Formicidae)

Summary There is high within-nest relatedness for functional queens (with corpora lutea), nonfunctional queens (without corpora lutea), and workers in polygynous nests of Leptothorax acervorum. The high functional queen relatedness suggests that young mated queens are adopted back to their mother nest. Functional queen relatedness does not change with the number of queens present in the nest, suggesting that the number of generations of queens, on average two to three, is rather stable. Worker relatedness decreases with increasing number of functional queens per nest (Tables 5, 6). The number of queens contributing offspring to the nest (mothers), estimated from worker and functional queen relatedness, is lower than the number of functional queens, particularly in highly polygynous nests. Estimates of number of mothers in monogynous nests indicate that these nests previously were polygynous (Table 7). There is no correlation between nest relatedness and distance between nests, and budding-off, if present, thus appears to be a rare mode of nest founding (Table 8). There are no indications of inbreeding in the two populations studied since the frequency of heterozygotes is as high as expected from random mating (Table 4). Most likely, polygyny is the rule in L. acervorum and serves to secure the presence of queens in the nest.     

Trail and recruitment pheromones in Camponotus socius (Hymenoptera: Formicidae)

Summary. Worker ants of Camponotus socius employ multicomponent signals during group recruitment to food sources and nest emigrations. The chemical signals consist of trail orientation pheromones that originate from the hindgut. Two components that elicit trail orientation behavior were identified: (2S,4R,5S)-2,4-dimethyl-5-hexanolide and 2,3-dihydro-3,5-dihydroxy-6-methylpyran-4-one. Although both compounds release trail following behavior, in choice tests the former compound is preferred significantly. The major recruitment signal appears to be formic acid discharged from the poison gland. Compounds identified in the Dufour gland secretions do not seem to be involved in the recruitment process.     

The social behavior of Leptothorax allardycei (Hymenoptera,Formicidae): time budgets and the evolution of worker reproduction

Summary The use of time by workers of the ant Leptothorax allardycei (Mann) is examined. Theoretical predictions are developed concerning the maximal lowering in colony reproductive output that is consistent with the evolution of worker production. Measurements are made of the effect of aggression on colony efficiency. Analysis of time budgets indicate that a typical ant spends a large fraction (0.55) of its time quiescent and another large fraction of time (0.32) involved in undifferentiated activity. Dominance activity and brood care together make up about 11% of the total time. The amount of time spent on dominance activity is negatively related to the amount of time spent on brood care, but positively related to the amount of time that an ant is active. The amount of time that an ant has available for brood care which is actually spent on brood care declines with the amount of time spent on dominance activity. The amount of time that a worker spends feeding liquid food to larvae is a function of hierarchy rank; alpha spends the most time, beta less and gamma still less. The spread of the trait of worker reproduction is examined theoretically, with particular regard to the associated costs to colony reproduction. Worker reproduction can spread through a population, under a variety of formulations, provided the cost to colony reproduction is less than some critical value in the neighborhood of 0.17–0.22 of the total colony output. The cost of worker reproduction in L. allardycei is estimated in two ways: as a time cost and as a reduction in the total number of brood tended per unit time. The two estimates of cost are 0.15 and 0.13 respectively. The reproductive options of the worker caste and the division of reproductive labor vary considerably between species. Reproduction by workers yields fitness differences between workers and results in competition among workers with the result that colony efficiency is affected.     

Trail discrimination signal of Lasius japonicus (Hymenoptera: Formicidae)

Summary. Trail-following behavior of Lasius japonicus was colony-specific in the field, while trail pheromone activity was not. We found that the footprint substance caused colony-specific trail-following behavior only when working in conjunction with the trail pheromone. The footprint substance alone did not lead the workers to follow trails. The substance consisted mainly of hydrocarbons with composition almost identical to that of cuticular hydrocarbons, except for the absence of n-alkanes. Nestmate workers shared footprint hydrocarbon profiles as well as cuticular hydrocarbons, but the profiles differed among colonies. We therefore consider that the footprint hydrocarbon profiles serve as the trail discrimination signal in L. japonicus.     

Caste and division of labor in leaf-cutter ants (Hymenoptera: Formicidae: Atta)

Summary Leaf cutting was selected for an evaluation of ergonomic efficiency in the fungus-growing ant Atta sexdens because it is performed largely by medias (head width 1.8–2.8 mm), which attend to relatively few other functions and hence are less likely to be evolutionarily compromised by the demands of competing tasks (Fig. 1).Three alternative a priori criteria of evolutionary optimization were envisioned that are consistent with natura selection theory: the reduction of predation by means of defense and evasion during foraging, the minimization of foraging time through skill and running velocity during foraging, and energetic efficiency, which must be evaluated with reference to both the energetic construction costs of new workers and the energetic cost of maintenance of the existing worker force.In order to measure the performance of various size groups within the A. sexdens worker caste in isolation, I devised the pseudomutant technique: in each experiment, groups of foraging workers were thinned out until only individuals of one size class were left outside the nest. Measurements were then made of the rate of attraction, initiative in cutting, and performance of each size group at head-width intervals of 0.4 mm (Figs. 2, 3, and 7). Other needed measurements were made in body weight, oxygen consumption, and running velocity (Figs. 5, 6, and 8).The size-frequency distribution ff leaf cutters in the A. sexdens conforms closely to the optimum predicted by the energetic efficiency criterion for harder forms of vegetation, such as rhododendron leaves. The distribution is optimum with reference to both construction and maintenance costs. The difference between the predicted and actual modal size groups specializing on leaf cutting is 10% or less of the total size range of the sexdens worker caste.A model was next constructed in which attraction and initiative were allowed to evolve genetically to uniform maximum levels. The theoretical maximum efficiency levels obtained by this means were found to reside in the head-width 2.6–2.8 mm size class, or 8% from the actual maximally efficiency class (head width 2.2–2.4 mm). In the activity of leaf cutting, A. sexdens can therefore be said to be not only at an adaptive optimum but also, within at most a relatively narrow margin of error, to have been optimized in the course of evolution.     

Worker size variability and foraging efficiency in Veromessor pergandei (Hymenoptera: Formicidae)

Summary As foragers of the harvester ant, Veromessor pergandei, travel further from their nest they spend significantly more time sampling seeds in experimental patches. Although accepted seeds are heavier than offered seeds, mass of accepted seed is not correlated with sampling time. Variably sized V. pergandei workers do not size-match; little, if any, variance in size of seed selected can be attributed to body size of forager. The lack of size-matching in V. pergandei suggests individual performance may be an inadequate measure of colony foraging success.     

Predatory behavior and chemical communication in two Metapone species (Hymenoptera:Formicidae)

Summary. Colonies of two species of Metapone (M. madagascarica, M. new species.) were collected in Madagascar and established in laboratory nests. It could be demonstrated that both species are specialist predators of termites (Cryptotermes kirbyi). During hunting the ants sting the termites and thereby paralyze and preserve the prey alive. In this way prey can be stored in the ant nest for extended periods. During foraging and colony emigrations the ants lay chemical trails with poison gland secretions. Among the seven compounds identified in the venom only methyl pyrrole-2-carboxylate elicits trail following behavior in both Metapone species. Received 11 February 2002, accepted 23 February 2002.     

Annual cycles in worker size of the seed-harvester antVeromessor pergandei (Hymenoptera: Formicidae)

Summary The seed-harvester antVeromessor pergandei Mayr is primitively polymorphic; workers are monophasically allometric. There is a distinct annual cycle in mean worker body size that replicates across colonies and habitats (Fig. 1); this cycle occurs through alteration of the worker size distribution (Fig. 2). There is little, if any, morphspecific task specialization by workers suggesting worker size variance is a colony-level adaptation permitting maintenance of a large and constant worker force during periods of resource fluctuation. Smaller workers appear in the foraging force following the triple crunch of reduced seed availability, reduced favorable times to forage, and alate production during winter months. Adult and startingV. pergandei colonies exhibit strong intraspecific territoriality, suggesting the selective advantage for maintenance of a large and constant worker force. Such selective pressures may have provided the initial variance in worker size distributions that led subsequently to specialized castespecific task performance in more distinctly polymorphic ant species.     

Queen dimorphism and reproductive strategies in the fire ant Solenopsis geminata (Hymenoptera: Formicidae)

Alate trapping studies of a monogyne population of the fire ant Solenopsis geminata indicate that two sizes of gynes are produced. Macrogynes, which participate in late spring and summer mating flights, are larger, fattier, and more than twice as heavy as microgynes, which participate in fall mating flights. Three patterns of gyne production were observed in 51 colonies studied: 35 produced macrogynes only, 9 produced microgynes only, and 7 produced both morphs, contributing to both summer and fall mating flights. Behavioral evidence and rearing studies suggest that macrogynes found new colonies independently, whereas microgynes achieve colony queen status by infiltrating or being adopted by established colonies. Of the total number of female alates collected from the trapped colonies, 56% were microgynes. However, because of their smaller size and lower fat content, microgynes made up only one-third of the caloric investment in female alates. By measuring the thorax lengths of queens from mature colonies, we determined that at least 56% were macrogynes and 35% or more were microgynes. These results indicate that as a reproductive strategy, colony investment in microgyne production may have at least as high a payoff as investment in macrogyne production.This is publication #24 of the Fire Ant Research Team     

Genetic relatedness of worker nestmates in Myrmica ruginodis (Hymenoptera : Formicidae) populations

Summary Genetic relatedness and mating structure of the red ant, Myrmica ruginodis, were studied from a large data set in several natural populations, one of which was a substructured archipelago population. Within-colony relatedness of worker nestmates was measured as genotypic correlation and mating structure as deviation from random mating; both were calculated from the genotype frequency data. The average relatedness of worker nestmates was rather high across populations (0.43–0.73), and random mating was the rule within populations. The data were used to deduce the social structure of populations. With one possible exception, populations turned out to be weakly polygynous, implying that they consisted of the macrogyne form of the species. The exceptional population was relatively more polygynous and inbred, suggesting that it might include the microgyne form. This exceptional population lived in a habitat not noticeably different from that of the other populations, challenging previous suggestions of the habitat specialization of the two forms. The colonies were facultatively polygynous, and a comparison with earlier studies shows that polygyny is functional. Relatedness of worker nestmates varied slightly among populations, indicating that the social structure of Myrmica colonies is flexible.     

Dominance orders in the ponerine ant Pachycondyla apicalis (Hymenoptera,Formicidae)

Summary Intracolony aggression among workers of the neotropical ponerine ant species Pachycondyla apicalis leads to dominance orders. Antagonistic interactions can entail either overt physical attacks with the subordinate individual often exhibiting a submissive posture or the robbing and destruction of eggs laid by nestmates. The single queen, however, was never observed either attacking or being attacked by any colony member. The hierarchical structure among workers consists of one dominant individual and several subordinates; the relationships among subordinate workers are unclear, however. We report for the first time a natural (nonmanipulated) change in the social status of individuals within an ant dominance order. Dominant workers usually had better developed ovaries, laid more eggs and were more frequently observed attending the egg pile than subordinate individuals. This pattern became even more striking when the queen was excluded from the colony. These results indicate that workers of P. apicalis lay eggs even in the presence of the queen. It is possible that some of these haploid eggs may develop into males.Offprint requests to: B. Holldobler     

Differential reproduction in multiple-queen colonies of the fire ant Solenopsis invicta (Hymenoptera: Formicidae)

Summary The contribution to maternity of workers and female sexuals over time by queens in six multiple-queen laboratory colonies of Solenopsis invicta was directly assessed by use of enzyme genetic markers. Queens contributed more equally to the worker pool than to the pool of sexuals in virtually all samples (Fig. 1), and individuals producing a substantial proportion of the workers often had low or no representation of their daughters in the pool of sexuals. Signficant disparity among queens in their relative production of sexual daughters was often evident, with dominance in production of sexuals by a given queen commonly occurring in association with a pronounced loss of weight followed shortly by her death. The results suggest that significant variability in short-as well as long-term reproductive success may occur among the distantly related queens associating in natural polygyne S. invicta nests. Variance in apportionment of maternity of sexuals did not appear to be simply related to varying levels of fecundity, suggesting that the common presumption that reproductive success can be equated with fecundity in polygyne social Hymenoptera may not be well founded. The observed variance also did not appear to result from a simple mechanism of kin recognition and discrimination by workers in the process of brood rearing. Rather, this variance may have largely resulted from either, 1) recognition of certain queens and their progeny coupled with preferential sexualization of these immatures by nurse workers, or, 2) queen biasing of eggs toward development as sexuals. The frequent association of weight loss and death of mother queens with high levels of sexual daughter production may be best explained by the latter mechanism.     

Colonial recognition of fungus in the fungus-growing ant Acromyrmex subterraneus subterraneus (Hymenoptera: Formicidae)

Summary. Leaf cutting ants live in symbiosis with a basidiomycete fungus that is exploited as a source of nutrients for the ant larvae. Tests of fungus transport demonstrated that Acromyrmex subterraneus subterraneus workers discriminate concolonial fungus from alien fungus, and rejected the latter. Larvae and pupae of the ant were used as controls. Chemical analysis of the fungus revealed a great similarity between its hydrocarbon profile and that found on the ant brood. Experiments with lures showed that chemical extracts from the fungus are responsible for this discrimination process. Moreover, the presence of brood inside the fungus seemed to be important for discrimination of the fungus by workers. Resident workers accepted concolonial broodless fungus less than concolonial fungus inoculated with brood odor. Fungus seems to acquire colonial odor passively, simply by contact with the brood. The impact of fungus volume present in the nest on closure of the colony is discussed. We show here for the first time the importance of a symbiotic vegetal organism in colonial recognition in social insects. Received 14 April 2000; accepted 29 September 2000     

The chemical basis for trail recognition in Lasius nipponensis (Hymenoptera: Formicidae)

Summary. In this paper we report on the chemical basis for trail recognition in Lasius nipponensis. On and near trails in the field, workers became aggressive against conspecific intruders and succeeded in protective contests, while intruders usually avoided confrontation. Such asymmetric interactions were also observed in the laboratory when two non-nestmate workers were sequentially placed in a clean glass dish, while mutual aggression was observed when they were placed simultaneously. Asymmetric aggression was also observed when the workers were placed in a dish previously conditioned by other workers, or when the dish was treated with the hexane rinse of the conditioned dish. The rinse contained a series of hydrocarbons, in which components and proportion were almost identical to those of the cuticular hydrocarbons of the workers, except for a lack of n-alkanes. Amounts of the hydrocarbons that remained on the dish were very small when all the tarsi of the workers were incapacitated. We therefore consider that the hydrocarbons are not adhesion of the cuticular hydrocarbons but secretion from the tarsi of workers, and serve as a key signal for trail recognition in this ant.