Dominique Fresneau

Individuality and colonial identity in ants: the emergence of the social representation concept

Colonial identity in social insects is based on nestmate recognition which is mediated through cuticular substances. Although this is considered to be distinct from kin recognition, it is possible that through evolution the signal mediating kinship was replaced by the signal mediating “nestmateship”. Cuticular hydrocarbons in Cataglyphis niger are responsible for modifying the ant’s aggressive behavior and are considered to have a similar function in other ants species. In ants, the postpharyngeal gland (PPG) serves as a storage organ for these cues and functions as a “gestalt” organ, with the gestalt being permanently updated. Its content is constantly being exchanged with nestmates through trophallaxis and allogrooming. We hypothesize that already in the primitive ponerine ants the PPG evolved as a gestalt organ even without trophallaxis. We discuss two alternative primary selective pressures for the evolution of trophallaxis: facilitating food exchange versus exchanging recognition cues. Callow workers seem to be characterized by a “cuticular chemical insignificance” followed by a “chemical integration” period when they acquire the gestalt of the colony and learn the associated template. We hypothesize that the template has evolved from a simple personal chemical reference in primitive species with small colonies to an internal representation of the colonial identity in larger colonies.

Formation of colony odor in ponerine ant Pachycondyla apicalis

The mechanisms operating in the formation and maintenance of colony odor in the ponerine ant Pachycondyla apicalis were studied using radioactive tracers. Using [l-14C]acetate as a precursor, the de novo biosynthesis and distribution of pentane-extractable lipids within the ants body were followed. Twenty-four hours after injection, newly synthesized alkanes, alkenes, as well as more polar lipids were found in the postpharyngeal gland, the epicuticle, and internally. Ants that had their mouths blocked with beeswax possessed significantly less radiolabeled lipids (all classes measured) in the postpharyngeal gland but had augmented levels in the internal pool and the epicuticle. Both hydrocarbons and more polar lipids were exchanged between the respective postpharyngeal glands and the epicuticle. The transfer to postpharyngeal glands was highest when both ants had an open mouth or when the recipient ant had an open mouth but the donors mouth was blocked. This suggests that the transfer to the postpharyngeal gland in this species is by allogrooming and not by trophallaxis. Transfer to the cuticle was low and comparable in all treatments. Behavioral observations during the first 6 hr of the dyadic encounters and in intact colonies confirmed that the ants did not engage in trophallaxis. The level of transfer to the postpharyngeal gland in P. apicalis was significantly lower than in representatives of other Formicidae subfamilies studied so far. We attribute this difference to the evolution of trophallaxis in the higher Formicidae.

Worldwide invasion by the little fire ant: routes of introduction and eco-evolutionary pathways

Biological invasions are generally thought to occur after human aided migration to a new range. However, human activities prior to migration may also play a role. We studied here the evolutionary genetics of introduced populations of the invasive ant Wasmannia auropunctata at a worldwide scale. Using microsatellite markers, we reconstructed the main routes of introduction of the species. We found three main routes of introduction, each of them strongly associated to human history and trading routes. We also demonstrate the overwhelming occurrence of male and female clonality in introduced populations of W. auropunctata, and suggest that this particular reproduction system is under selection in human‐modified habitats. Together with previous researches focused on native populations, our results suggest that invasive clonal populations may have evolved within human modified habitats in the native range, and spread further from there. The evolutionarily most parsimonious scenario for the emergence of invasive populations of the little fire ant might thus be a two‐step process. The W. auropunctata case illustrates the central role of humans in biological change, not only due to changes in migration patterns, but also in selective pressures over species.

A Basis for Spatial and Social Patterns in Ant Species: Dynamics and Mechanisms of Aggregation

Aggregation is usually studied in functional terms, forgetting mechanisms. In this paper, experimental results on the ant Lasius niger, complemented by a model, allow us to understand the mechanisms responsible for aggregation and to study the influence of the population density on this phenomenon. The results show (1) a high level of aggregation and the emergence of a large cluster; (2) that aggregation results from an amplification mechanism—the greater the number of ants inside a cluster, the greater the time spent by one ant in this cluster; and (3) that population density has only a weak influence on the aggregation process. This method of analysis and these results can certainly be extended not only in social insects but also in other species, like subsocial arthropods.

Dynamics of aggregation in Lasius niger (Formicidae): influence of polyethism

SummaryPolyethism is a well-known phenomenon in social insects. How this phenomenon influences interactions among individuals, the spatial distribution in the nest is, on the other hand, very rarely documented. Therefore, we conducted experiments on the ant Lasius niger to observe the influence of polyethism on aggregation, by distinguishing two groups of ants: the brood-tenders and the foragers. We show a great difference in their self aggregation level. Brood-tenders are characterized by a rapid and dense gathering in one main stable cluster while foragers gather in several small unstable clusters. We show experimentally and verify with a model that this difference in behaviour is based on a smaller probability of leaving a cluster for the brood-tenders. Aggregation in the mixed case (groups composed of brood-tenders and foragers) is very close to that of the pure forager case, showing a decrease in the level of aggregation of the brood-tenders respecting to the pure group of brood-tenders. Nevertheless, experimental results supported by the results of the model, show that ants do not change their own behaviour when the two groups are together. Therefore, the decrease of the aggregation of brood-tenders in the mixed case can be explained by a difference in the dynamics between brood-tenders and foragers.

Hydrocarbon distribution and colony odour homogenisation in Pachycondyla apicalis

SummaryWithin and between individuals hydrocarbon (HC)-circulation was studied in Pachycondyla apicalis workers, using radioactive labeling. Newly synthesized HCs occurred both in the PPG and on the epicuticle in appreciable amounts, lesser quantities were found in the crop. The front basitarsal brush contained a greater amount of radiolabeled HCs than could be predicted from its surface area, suggesting preferential secretion to these organs. We propose that the newly synthesized HCs are secreted primarily to the front basitarsal brushes and are thereafter either distributed throughout the body surface, or cleared via the PPG and the alimentary canal.Using labeled HCs as a model, we tracked the time-dependent dispersion of cuticular lipids among 11 workers, one of which was prelabeled for 24 hours. Distribution among the recipients became progressively uniform, reaching near homogenization between 5–10 days. The mean HCs transfer of P. apicalis to the PPG was substantially lower compared to that of Camponotus fellah or Aphaenogaster senilis. In contrast, transfer to the cuticle in this species was superior. We attribute the low transfer to the PPG to the inefficacy of passive body contact characteristic of P. apicalis, as opposed to trophallaxis and/or allogrooming that typify the other two species. The higher occurrence of radiolabeled HCs in P. apicalis cuticle can be attributed to their accumulation in the basitarsal brushes. The impact of cuticular lipid transfer and formation of uniform colony odour, as opposed to the maintenance of an idiosyncratic caste-specific composition, are discussed.

Facultative polygyny in Ectatomma tuberculatum (Formicidae, Ectatomminae)

Summary.Polygyny, the presence of several mated queens within the same colony, is widespread in insect societies. This phenomenon is commonly associated with ecological constraints such as limited nest sites. In habitats where solitary nest foundation is risky, monogynous colonies can reintegrate young daughter queens (secondary polygyny). We studied the reproductive structure (i.e. queen number) of the ectatommine ant Ectatomma tuberculatum from Bahia State, Brazil. This species was found to present facultative polygyny: out of a total of 130 colonies collected, 39.2% were monogynous, while 43.8% were polygynous. Polygynous colonies had significantly more workers than monogynous ones. Queen number in polygynous colonies ranged from 2 to 26, with an average of 4 ± 4 queens per colony. All nestmate queens were egg-layers with no apparent dominance hierarchy or agonistic behavior. Non-nestmate queens were adopted by monogynous colonies suggesting that polygyny is secondary, originating through queen adoption. This species is characterized by an open recognition system, which probably allows a switch from monogynous to polygynous colonies. The behavioral acts of queens showed that resident queens remained frequently immobile on or near the brood, contrarily to alien or adopted queens and gynes. In addition, monogynous queens showed no behavioral or physiological (i.e. by ovarian status) differences in comparison with polygynous ones. Secondary or facultative polygyny, probably associated with queen adoption, may have been favored in particular environmental conditions. Indeed, by increasing colony productivity (i.e. number of workers) and territory size (by budding and polydomy), polygyny could uphold E. tuberculatum as a dominant species in the mosaic of arboreal ants in Neotropical habitats.

A study of polyethism in a ponerine ant: Neoponera apicalis (Hymenoptera, formicidae)

Abstract The division of labour in a colony of Neoponera apicalis Latr. was studied, observations being made over a period of 21 days during which each member of the society was individually monitored. Processing the results by correlation analysis made it possible to establish the relationships between the various behavioural patterns observed. These were organized around three focal activities: care of the brood, non-specific activities in the nest and foraging. The division of ants into these three categories was relatively consistent. The social status of individual ants was determined by cluster analysis. This revealed the existence of two further subgroups: domestic workers and pre-foragers. The age of individuals is a probable factor governing the division of labour. This organization is consistent with that observed in higher species. However, the participation of winged females, and probably queens, in various tasks on the same basis as the workers may be considered a primitive trait characteristic of species of ponerine ants.

How an Ant Manages to Display Individual and Colonial Signals by Using the Same Channel

Cuticular hydrocarbons are used by some ants to discriminate nestmates from nonnestmates. Every member of the colony bears the same pattern because they are continuously exchanged among nestmates. The postpharyngeal gland (PPG) stores the blend of hydrocarbons and is involved in the distribution of this common mixture. However, some individuals might display individual information on the cuticle (such as a chemical signal of fertility) that must not be mixed within the common pool. We investigated how this paradox is solved in the ant Pachycondyla goeldii by analyzing the nature and localization of colonial and fertility signals. Workers in a queenless condition showed a dominance hierarchy that was correlated with ovarian development. Hydrocarbons from the cuticle and the PPG analyzed by gas chromatography (GC) and identified by GC-mass spectrometry showed a clear discrimination among colonies, supporting the involvement of the PPG in the colonial identity signal. We identified and selected 11 cuticular hydrocarbons that permitted us to discriminate ovarian development classes and that might function as a fertility signal. They allowed clear colony discrimination as well, which suggests that the two signals (the individual signal of fertility and the common signal of colony identity) can be conveyed by the same compounds. However, the hydrocarbons in the PPG did not discriminate among ovarian developmental classes, suggesting that the portion of variation in the cuticular hydrocarbons constituting the fertility signal is superimposed on the signal of colony identity.

Postmating changes in cuticular chemistry and visual appearance in Ectatomma tuberculatum queens (Formicidae: Ectatomminae)

In the ectatommine ant Ectatomma tuberculatum, the visual appearance of queens changes after mating and ovarian development in that their cuticle turns from shiny to matte. In this study, we have shown that this change seems to be caused by 15-fold accumulation of hydrocarbons, in particular heptacosane that covers the multiple grooves present on the cuticular surface creating a wax coat in mated fully fertile queens. Analyses of the scrapped wax revealed that it is composed largely of heptacosane. Peak-by-peak comparison of the cuticular hydrocarbon (CHC) composition of mated, virgin with developed ovaries and virgin with nondeveloped ovaries revealed significant differences between the queen groups. Although the total amount of the CHC of virgin queens with developed ovaries was not higher than virgin queens that did not have developed ovaries, the composition showed a shift toward the mated queen. While it is possible that the large accumulation of hydrocarbons may give extra physical and chemical protection to queens, we propose that the switch in the relative abundance of heptacosane and nonacosane and perhaps of other components is indicative of being a mating and fertility cue. This is the first report in social insects where external chemical changes are accompanied by changes in visual appearance.