Queen movement during colony emigration in the facultatively polygynous ant Pachycondyla obscuricornis

In ants, nest relocations are frequent but nevertheless perilous, especially for the reproductive caste. During emigrations, queens are exposed to predation and face the risk of becoming lost. Therefore the optimal strategy should be to move the queen(s) swiftly to a better location, while maintaining maximum worker protection at all times in the new and old nests. The timing of that event is a crucial strategic issue for the colony and may depend on queen number. In monogynous colonies, the queen is vital for colony survival, whereas in polygynous colonies a queen is less essential, if not dispensable. We tested the null hypothesis that queen movement occurs at random within the sequence of emigration events in both monogynous and polygynous colonies of the ponerine ant Pachycondyla obscuricornis. Our study, based on 16 monogynous and 16 polygynous colony emigrations, demonstrates for the first time that regardless of the number of queens per colony, the emigration serial number of a queen occurs in the middle of all emigration events and adult ant emigration events, but not during brood transport events. It therefore appears that the number of workers in both nests plays an essential role in the timing of queen movement. Our results correspond to a robust colony-level strategy since queen emigration is related neither to colony size nor to queen number. Such an optimal strategy is characteristic of ant societies working as highly integrated units and represents a new instance of group-level adaptive behaviors in social insect colonies.     

Queen regulates biogenic amine level and nestmate recognition in workers of the fire ant, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>

Nestmate recognition is a critical element in social insect organization, providing a means to maintain territoriality and close the colony to parasites and predators. Ants detect the colony chemical label via their antennae and respond to the label mismatch of an intruder with aggressive behavior. In the fire ant, Solenopsis invicta, worker ability to recognize conspecific nonnestmates decreases if the colony queen is removed, such that they do not recognize conspecific nonnestmates as different. Here, we tested the hypothesis that the presence of the colony queen influences the concentration of octopamine, a neuromodulator, in worker ants, which in turn has an effect on nestmate recognition acuity in workers. We demonstrate that queenless workers exhibit reduced brain octopamine levels and reduced discriminatory acuteness; however, feeding queenless workers octopamine restored both. Dopamine levels are influenced by honeybee queen pheromones; however, levels of this biogenic amine were unchanged in our experiments. This is the first demonstration of a link between the presence of the colony queen, a worker biogenic amine, and conspecific nestmate recognition, a powerful expression of colony cohesion and territoriality.     

A colony-level response to disease control in a leaf-cutting ant

Parasites and pathogens often impose significant costs on their hosts. This is particularly true for social organisms, where the genetic structure of groups and the accumulation of contaminated waste facilitate disease transmission. In response, hosts have evolved many mechanisms of defence against parasites. Here we present evidence that Atta colombica, a leaf-cutting ant, may combat Escovopsis, a dangerous parasite of Atta's garden fungus, through a colony-level behavioural response. In A. colombica, garden waste is removed from within the colony and transported to the midden - an external waste dump - where it is processed by a group of midden workers. We found that colonies infected with Escovopsis have higher numbers of workers on the midden, where Escovopsis is deposited. Further, midden workers are highly effective in dispersing newly deposited waste away from the dumping site. Thus, the colony-level task allocation strategies of the Atta superorganism may change in response to the threat of disease to a third, essential party.     

Fire ant venom alkaloids act as key attractants for the parasitic phorid fly, Pseudacteon tricuspis (Diptera: Phoridae)

The phorid fly, Pseudacteon tricuspis Borgmeier, is an introduced parasitoid of imported fire ants, Solenopsis spp., in the USA. Although the assumption that phorid flies use fire ant alarm pheromones for host location is probably true, we demonstrated in a previous study the possible involvement of other ant semiochemicals in the response of P. tricuspis to fire ants. This study was conducted to determine the glandular sources and identity of the semiochemicals mediating this interaction. First, we tested the electroantennogram response of P. tricuspis to extracts of key body parts and glands of workers of the red imported fire ant, S. invicta Buren. The results confirm that the poison (venom) gland/sac is the key source of compounds which elicited strong antennal activity in P. tricuspis. Follow-up studies were conducted by using a combination of bioassay-guided fractionation and behavioral bioassays to test the hypothesis that attraction of this parasitoid to fire ants is mediated by venom alkaloids. The results confirm the response of P. tricuspis to physiologically relevant amounts of the two venom alkaloid fractions (cis and trans alkaloid fractions) of S. invicta. Further analysis by coupled gas chromatography–electroantennogram detection revealed nine venom alkaloid components including two novel 2,6-dialkylpiperideines that elicited significant antennal activity in P. tricuspis. This is the first demonstration of the role of venom alkaloids of ants as attractants for their natural enemies. We propose a semiochemical-mediated host location mechanism for P. tricuspis involving both alarm pheromones and venom alkaloids. The ecological significance of these findings, including the attraction of male P. tricuspis to fire ant venom alkaloids, possibly for mate location, is discussed.     

Non-specific association between filamentous bacteria and fungus-growing ants

Fungus-growing ants and their fungal cultivar form a highly evolved mutualism that is negatively affected by the specialized parasitic fungus Escovopsis. Filamentous Pseudonocardia bacteria occurring on the cuticle of attine ants have been proposed to form a mutualistic interaction with these ants in which they are vertically transmitted (i.e. from parent to offspring colonies). Given a strictly vertical transmission of Pseudonocardia, the evolutionary theory predicts a reduced genetic variability of symbionts among ant lineages. The aim of this study was to verify whether actinomycetes, which occur on Acromyrmex octospinosus leaf-cutting ants, meet this expectation by comparing their genotypic variability with restriction fragment length polymorphisms. Multiple actinomycete strains could be isolated from both individual ant workers and colonies (one to seven strains per colony). The colony specificity of actinomycete communities was high: Only 15% of all strains were isolated from more than one colony, and just 5% were present in both populations investigated. Partial sequencing of 16S ribosomal deoxyribonucleic acid of two of the isolated strains assigned both of them to the genus Streptomyces. Actinomycetes could also be isolated from workers of the two non-attine ant species Myrmica rugulosa and Lasius flavus. Sixty-two percent of the strains derived from attine ants and 80% of the strains isolated from non-attine ants inhibited the growth of Escovopsis. Our data suggest that the association between attine ants and their actinomycete symbionts is less specific then previously thought. Soil-dwelling actinomycetes may have been dynamically recruited from the environment (horizontal transmission), probably reflecting an adaptation to a diverse community of microbial pathogens.     

Effect of time on colony odour stability in the ant <Emphasis Type="Italic">Formica exsecta</Emphasis>

Among social insects, maintaining a distinct colony profile allows individuals to distinguish easily between nest mates and non-nest mates. In ants, colony-specific profiles can be encoded within their cuticular hydrocarbons, and these are influenced by both environmental and genetic factors. Using nine monogynous Formica exsecta ant colonies, we studied the stability of their colony-specific profiles at eight time points over a 4-year period. We found no significant directional change in any colony profile, suggesting that genetic factors are maintaining this stability. However, there were significant short-term effects of season that affected all colony profiles in the same direction. Despite these temporal changes, no significant change in the profile variation within colonies was detected: each colony’s profile responded in similar manner between seasons, with nest mates maintaining closely similar profiles, distinct from other colonies. These findings imply that genetic factors may help maintain the long-term stability of colony profile, but environmental factors can influence the profiles over shorter time periods. However, environmental factors do not contribute significantly to the maintenance of diversity among colonies, since all colonies were affected in a similar way.     

The role of tending ants in host plant selection and egg parasitism of two facultative myrmecophilous butterflies

Ovipositing adult females of myrmecophilous lycaenids are expected to select plants based on ant presence in order to maximize the survivorship of immature stages. Usually, larvae feed ants with honey-like solutions and, in turn, ants ward off parasitoids. Nonetheless, a rarely investigated approach is whether ant partners can also extend their protective behavior towards lycaenids eggs. Here, we investigated the ant-related oviposition pattern of Allosmaitia strophius and Rekoa marius; then, we compared egg parasitism according to the presence of ants. Lycaenid oviposition and egg parasitism (in percent) were experimentally compared in ant-present and ant-excluded treatments. The study plant, Heteropterys byrsonimifolia, is an extrafloral nectaried shrub which supports several ant species. We sampled 280 eggs, of which 39.65 % belonged to A. strophius and 60.35 % to R. marius. Both lycaenids eggs were significantly more abundant on branches with ants, especially those with Camponotus crassus and Camponotus blandus, two ant species known to attend to lycaenids. A. strophius and R. marius parasitism was 4.5- and 2.4-fold higher, respectively, in ant-present treatments, but the results were not statistically significant. Our study shows that ant-mediated host plant selection in lycaenids might be much more widespread than previously thought, and not restricted to obligate myrmecophilous species. Tending ants may be inefficient bodyguards of lycaenid eggs, because unlike larvae which release sugared liquids, eggs do not offer obvious rewards to ants. Ants can ward off parasitoids of larvae, as observed elsewhere, but our findings show that positive ant–lycaenid interactions are conditional and depend on immature ontogeny.     

Clonal structure affects the assembling behavior in the Japanese queenless ant <Emphasis Type="Italic">Pristomyrmex punctatus</Emphasis>

The queenless ant Pristomyrmex punctatus (Hymenoptera: Myrmicinae) has a unique society that differs from those of other typical ants. This species does not have a queen, and the workers lay eggs and produce their clones parthenogenetically. However, a colony of these ants does not always comprise members derived from a single clonal line. In this study, we examined whether P. punctatus changes its “assembling behavior” based on colony genetic structure. We prepared two subcolonies—a larger one comprising 200 individuals and a smaller one comprising 100 individuals; these subcolonies were established from a single stock colony. We investigated whether these subcolonies assemble into a single nest. The genetically monomorphic subcolonies (single clonal line) always fused into a single nest; however, the genetically polymorphic subcolonies (multiple clonal lines) did not tend to form a single colony. The present study is the first to demonstrate that the colony genetic structure significantly affects social viscosity in social insects.     

Nest- and colony-mate recognition in polydomous colonies of meat ants (Iridomyrmex purpureus)

Workers of polydomous colonies of social insects must recognize not only colony-mates residing in the same nest but also those living in other nests. We investigated the impact of a decentralized colony structure on colony- and nestmate recognition in the polydomous Australian meat ant (Iridomyrmex purpureus). Field experiments showed that ants of colonies with many nests were less aggressive toward alien conspecifics than those of colonies with few nests. In addition, while meat ants were almost never aggressive toward nestmates, they were frequently aggressive when confronted with an individual from a different nest within the same colony. Our chemical analysis of the cuticular hydrocarbons of workers using a novel comprehensive two-dimensional gas chromatography technique that increases the number of quantifiable compounds revealed both colony- and nest-specific patterns. Combined, these data indicate an incomplete transfer of colony odor between the nests of polydomous meat ant colonies.     

Forest clearing and sex ratio in forest-dwelling wood ant <Emphasis Type="Italic">Formica aquilonia</Emphasis>

Sex ratios of ants have been shown to vary with food resource levels in several studies, but it is not known whether forest clear-cutting has any effect on sex ratio of aphid-tending forest-dwelling ants. We investigated whether the offspring sex ratio of the forest dwelling ant Formica aquilonia varied as a response to clear-cutting. We found that the proportion of males was smaller in clear-cuts than in adjacent forests. Our results are among the first showing that anthropogenic changes in forest structures may have a potential to modify sex ratios of social insects and other forest-dwelling animals.     

<Emphasis Type="Italic">Gnamptogenys hartmani</Emphasis> Wheeler (Ponerinae: Ectatommini): an agro-predator of<Emphasis Type="Italic"> Trachymyrmex</Emphasis> and<Emphasis Type="Italic"> Sericomyrmex</Emphasis> fungus-growing ants

The fungus gardens of fungus-growing ants are a potentially valuable resource for exploitation by natural enemies, but few of these antagonistic interactions have been studied. Here we describe key aspects of the behavioral ecology of Gnamptogenys hartmani (Ponerinae: Ectatommini), a specialized agro-predator of Trachymyrmex and Sericomyrmex fungus-growing ants in Panama. Raiding columns of G. hartmani attack and usurp nests with remarkably little effort: a few intruding workers are sufficient to cause panic among the attine ants and make them abscond from the nest. Both G. hartmani larvae and adults consume the fungus and the host brood, after which the colony migrates to a new fungus-growing ant nest discovered by scouting workers. The morphology of the G. hartmani larval mouthparts is similar to that of Gnamptogenys species with a non-fungal diet. However, we suggest that the presence of long spinules on the larval mandibles in the genus Gnamptogenys, comparable to those found in attine larvae, may have pre-adapted G. hartmani to fungus eating. G. hartmani workers do not actively maintain or modify fungus gardens, which makes them less efficient exploiters than Megalomyrmex, the only other agro-predatory ant species known so far.     

Nutrient compensatory foraging in a free-living social insect

The geometric framework model predicts that animal foraging decisions are influenced by their dietary history, with animals targeting a combination of essential nutrients through compensatory foraging. We provide experimental confirmation of nutrient-specific compensatory foraging in a natural, free-living population of social insects by supplementing their diet with sources of protein- or carbohydrate-rich food. Colonies of the ant Iridomyrmex suchieri were provided with feeders containing food rich in either carbohydrate or protein for 6 days, and were then provided with a feeder containing the same or different diet. The patterns of recruitment were consistent with the geometric framework: while feeders with a carbohydrate diet typically attracted more workers than did feeders with protein diet, the difference in recruitment between the two nutrients was smaller if the colonies had had prior access to carbohydrate than protein. Further, fewer ants visited feeders if the colony had had prior access to protein than to carbohydrates, suggesting that the larvae play a role in worker foraging behaviour.     

Colony size affects division of labour in the ponerine ant<Emphasis Type="Italic"> Rhytidoponera metallica</Emphasis>

In theory, larger colonies of social insects should have greater colony organisation. While inter-specific comparative studies provide support for this idea, there is little direct intra-specific evidence. We investigated differences in task specialisation between large (>450 workers) and small (<80 workers) colonies of the ponerine ant Rhytidoponera metallica. Observations of individually marked young or old workers revealed greater task specialisation in large colonies. Age polyethism was detected in large but not small colonies. In large colonies, old workers spent significantly more time foraging than young workers did, while young workers spent more time caring for brood. In small colonies, young and old workers spent a similar amount of time foraging and caring for brood. This difference in task allocation patterns in large and small colonies was associated with a difference in contact rates between workers. Workers in small colonies have a lower contact rate between nestmates and a greater variability in time between contacts than workers from large colonies.     

Size-correlated division of labour and spatial distribution of workers in the driver ant,<Emphasis Type="Italic"> Dorylus molestus</Emphasis>

Driver ants ( Dorylus spp.) show a high degree of worker polymorphism. Previous reports suggest that large Dorylus workers are specialised for defensive tasks. In this study, we first quantitatively tested whether there is a size-correlated division of defensive labour among workers. Second, we determined whether the spatial distribution of workers outside the nest can be predicted based on such size-specific differences in task allocation. We show that the division of defensive behaviour among different-sized workers is not strict. However, there is a significant correlation between worker size and the tendency to carry out defensive tasks. First, workers of larger size were more likely than smaller workers to participate in colony defence. Second, larger workers were more frequent near the nest containing the reproducing individuals and the brood. Finally, large workers were more common in open sections of the trail than in covered sections, which are likely to be less exposed to predators.     

Effect of post-fire resprouting on leaf fluctuating asymmetry, extrafloral nectar quality, and ant–plant–herbivore interactions

Fires in the Cerrado savanna are a severe form of disturbance, but some species are capable of resprouting afterwards. It is unknown, however, how and whether post-fire resprouting represents a stressful condition to plants and how their rapid re-growth influences both the production of biochemical compounds, and interactions with mutualistic ants. In this study, we examined the influence of post-fire resprouting on biotic interactions (ant–plant–herbivore relationships) and on plant stress. The study was performed on two groups of the extrafloral nectaried shrub Banisteriopsis campestris (Malpighiaceae); one group was recovering from fire while the other acted as control. With respect to biotic interactions, we examined whether resprouting influenced extrafloral nectar concentration (milligrams per microliter), the abundance of the ant Camponotus crassus and leaf herbivory rates. Plant stress was assessed via fluctuating asymmetry (FA) analysis, which refers to deviations from perfect symmetry in bilaterally symmetrical traits (e.g., leaves) and indicates whether species are under stress. Results revealed that FA, sugar concentration, and ant abundance were 51.7 %, 35.7 % and 21.7 % higher in resprouting plants. Furthermore, C. crassus was significantly associated with low herbivory rates, but only in resprouting plants. This study showed that post-fire resprouting induced high levels of plant stress and influenced extrafloral nectar quality and ant-herbivore relationships in B. campestris. Therefore, despite being a stressful condition to the plant, post-fire resprouting individuals had concentrated extrafloral nectar and sustained more ants, thus strengthening the outcomes of ant–plant mutualism.     

Biogenic amine levels, reproduction and social dominance in the queenless ant Streblognathus peetersi

Social harmony often relies on ritualised dominance interactions between society members, particularly in queenless ant societies, where colony members do not have developmentally predetermined castes but have to fight for their status in the reproductive and work hierarchy. In this behavioural plasticity, their social organisation resembles more that of vertebrates than that of the “classic” social insects. The present study investigates the neurochemistry of the queenless ant species, Streblognathus peetersi, to better understand the neural basis of the high behavioural plasticity observed in queenless ants. We report measurements of brain biogenic amines [octopamine, dopamine, serotonin] of S. peetersi ants; they reveal a new set of biogenic amine influences on social organisation with no common features with other “primitively organised societies” (bumble bees) and some common features with “highly eusocial” species (honey bees). This similarity to honey bees may either confirm the heritage of queenless species from their probably highly eusocial ancestors or highlight independent patterns of biogenic amine influences on the social organisation of these highly derived species.     

Circadian consequences of social organization in the ant species<Emphasis Type="Italic"> Camponotus compressus</Emphasis>

The locomotor activity rhythm of different castes of the ant species Camponotus compressus was monitored individually under laboratory light/dark (LD) cycles, and under continuous darkness (DD). The colony of this ant species comprises two sexual castes, the queens and the males, and three worker castes, namely the major, media, and minor workers. The virgin males and virgin queens display rhythmic activity patterns, but the mated queens were arrhythmic while laying eggs, with the rhythmicity resuming soon after egg-laying. Under the LD regime, major workers showed nocturnal patterns, while about 75% of the media workers displayed nocturnal patterns and about 25% showed diurnal patterns. Under the DD regime, most major workers exhibited circadian rhythm of activity with a single steady state, whereas media workers displayed two types of activity patterns, with activity patterns changing after 6–9 days in DD (turn-arounds). The pre-turn-around of the ants that showed nocturnal activity patterns during LD entrainment was <24 h after release into DD, which then became >24 h, after 6–9 days. On the other hand, the pre-turn-around of those ants that exhibited diurnal patterns during LD entrainment was first >24 h after release into DD, and then became <24 h, after 6–9 days. The activity of the minor workers neither entrained to LD cycles nor showed any sign of free-run in DD. It appears that the circadian clocks of the ant species C. compressus are flexible, and may perhaps depend upon the tasks assigned to them in the colony.     

Foraging ants trade off further for faster: use of natural bridges and trunk trail permanency in carpenter ants

Trail-making ants lay pheromones on the substrate to define paths between foraging areas and the nest. Combined with the chemistry of these pheromone trails and the physics of evaporation, trail-laying and trail-following behaviours provide ant colonies with the quickest routes to food. In relatively uniform environments, such as that provided in many laboratory studies of trail-making ants, the quickest route is also often the shortest route. Here, we show that carpenter ants (Camponotus rufipes), in natural conditions, are able to make use of apparent obstacles in their environment to assist in finding the fastest routes to food. These ants make extensive use of fallen branches, twigs and lianas as bridges to build their trails. These bridges make trails significantly longer than their straight line equivalents across the forest floor, but we estimate that ants spend less than half the time to reach the same point, due to increased carriage speed across the bridges. We also found that these trails, mainly composed of bridges, are maintained for months, so they can be characterized as trunk trails. We suggest that pheromone-based foraging trail networks in field conditions are likely to be structured by a range of potentially complex factors but that even then, speed remains the most important consideration.     

Winged queens replaced by reproductives smaller than workers in <Emphasis Type="Italic">Mystrium</Emphasis> ants

In ants, winged queens that are specialized for independent colony foundation can be replaced by wingless reproductives better adapted for colony fission. We studied this shift in reproductive strategy by comparing two Mystrium species from Madagascar using morphometry, allometry and dissections. Mystrium rogeri has a single dealate queen in each colony with a larger thorax than workers and similar mandibles that allow these queens to hunt during non-claustral foundation. In contrast, Mystrium ‘red’ lacks winged queens and half of the female adults belong to a wingless ‘intermorph’ caste smaller and allometrically distinct from the workers. Intermorphs have functional ovaries and spermatheca while those of workers are degenerate. Intermorphs care for brood and a few mate and reproduce making them an all-purpose caste that takes charge of both work and reproduction. However, their mandibles are reduced and inappropriate for hunting centipedes, unlike the workers’ mandibles. This together with their small thorax disallow them to perform independent colony foundation, and colonies reproduce by fission. M. rogeri workers have mandibles polymorphic in size and shape, which allow for all tasks from brood care to hunting. In M. ‘red’, colonial investment in reproduction has shifted from producing expensive winged queens to more numerous helpers. M. ‘red’ intermorphs are the first case of reproductives smaller than workers in ants and illustrate their potential to diversify their caste system for better colonial economy.