Larval memory affects adult nest-mate recognition in the ant Aphaenogaster senilis

Prenatal olfactory learning has been demonstrated in a wide variety of animals, where it affects development and behaviour. Young ants learn the chemical signature of their colony. This cue-learning process allows the formation of a template used for nest-mate recognition in order to distinguish alien individuals from nest-mates, thus ensuring that cooperation is directed towards group members and aliens are kept outside the colony. To date, no study has investigated the possible effect of cue learning during early developmental stages on adult nest-mate recognition. Here, we show that odour familiarization during preimaginal life affects recognition abilities of adult Aphaenogaster senilis ants, particularly when the familiarization process occurs during the first larval stages. Ants eclosed from larvae exposed to the odour of an adoptive colony showed reduced aggression towards familiar, adoptive individuals belonging to this colony compared with alien individuals (true unfamiliar), but they remained non-aggressive towards adult individuals of their natal colony. Moreover, we found that the chemical similarity between the colony of origin and the adoptive colony does not influence the degree of aggression, meaning that the observed effect is likely to be due only to preimaginal learning experience. These results help understanding the developmental processes underlying efficient recognition systems.     

The look of royalty: visual and odour signals of reproductive status in a paper wasp

Reproductive conflicts within animal societies occur when all females can potentially reproduce. In social insects, these conflicts are regulated largely by behaviour and chemical signalling. There is evidence that presence of signals, which provide direct information about the quality of the reproductive females would increase the fitness of all parties. In this study, we present an association between visual and chemical signals in the paper wasp Polistes satan. Our results showed that in nest-founding phase colonies, variation of visual signals is linked to relative fertility, while chemical signals are related to dominance status. In addition, experiments revealed that higher hierarchical positions were occupied by subordinates with distinct proportions of cuticular hydrocarbons and distinct visual marks. Therefore, these wasps present cues that convey reliable information of their reproductive status.     

Ants distinguish neighbors from strangers

Summary Ants are known to distinguish their own nests and nestmates from all others, using colony-specific odors. Here I show that harvester ants (Pogonomyrmex barbatus) can further distinguish between two kinds of non-nestmates of the same species: neighbors and strangers. Interactions between colonies were thought to depend on the numbers of alien ants that each colony encounters on its territory. The results described here show that such interactions also depend on information about colony identity. Encounters on foraging trails with ants from neighboring colonies, deter foraging more than encounters with ants from distant ones. The history of interactions between particular pairs of colonies may have important effects on intraspecific competition for food.     

Chemical basis of nest-mate discrimination in the ant Formica exsecta.

Distinguishing nest-mates from non-nest-mates underlies key animal behaviours, such as territoriality, altruism and the evolution of sociality. Despite its importance, there is very little empirical support for such a mechanism in nature. Here we provide data that the nest-mate recognition mechanism in an ant is based on a colony-specific Z9-alkene signature, proving that surface chemicals are indeed used in ant nest-mate recognition as was suggested 100 years ago. We investigated the cuticular hydrocarbon profiles of 10 Formica exsecta colonies that are composed almost entirely of a Z9-alkene and alkane component. Then we showed that worker aggression is only elicited by the Z9-alkene part. This was confirmed using synthetic Z9-alkene and alkane blends matched to the individual colony profiles of the two most different chemical colonies. In both colonies, only glass beads with 'nest-mate' alkene profiles received reduced aggression. Finally, changing the abundance of a single Z9-alkene on live ants was shown to significantly increase the aggression they received from nest-mates in all five colonies tested. Our data suggest that nest-mate discrimination in the social insects has evolved to rely upon highly sensitive responses to relatively few compounds.     

Nest-mate recognition in Manuelia postica (Apidae: Xylocopinae): an eusocial trait is present in a solitary bee

In eusocial Hymenoptera, females are more tolerant towards nest-mate than towards non-nest-mate females. In solitary Hymenoptera, females are generally aggressive towards any conspecific female. Field observations of the nest biology of Manuelia postica suggested nest-mate recognition. Experiments were performed involving two live interacting females or one live female interacting with a dead female. Live females from different nests were more intolerant to each other than females from the same nest. Females were more intolerant towards non-nest-mate than towards nest-mate dead females. When dead females were washed with pentane, no differences in tolerant and intolerant behaviours were detected between non-nest-mate and nest-mate females. Females were more intolerant towards nest-mate female carcasses coated with the cuticular extract from a non-nest-mate than towards non-nest-mate female carcasses coated with the cuticular extract from a nest-mate. The compositions of the cuticular extracts was more similar between females from the same nest than between females from different nests. The results demonstrate for the first time nest-mate recognition mediated by cuticular chemicals in a largely solitary species of Apidae. The position of Manuelia at the base of the Apidae phylogeny suggests that nest-mate recognition in eusocial species apical to Manuelia represents the retention of a primitive capacity in Apidae.     

Chemoreception: identifying friends and foes

The vomeronasal organ detects chemical cues that trigger sexual, aggressive and defensive behaviors. An in situ hybridization analysis has identified the specificities of nearly a hundred VNO receptors and elucidated the logic by which they encode these cues.     

Learning and discrimination of cuticular hydrocarbons in a social insect

Social insect cuticular hydrocarbon (CHC) mixtures are among the most complex chemical cues known and are important in nest-mate, caste and species recognition. Despite our growing knowledge of the nature of these cues, we have very little insight into how social insects actually perceive and discriminate among these chemicals. In this study, we use the newly developed technique of differential olfactory conditioning to pure, custom-designed synthetic colony odours to analyse signal discrimination in Argentine ants, Linepithema humile. Our results show that tri-methyl alkanes are more easily learned than single-methyl or straight-chain alkanes. In addition, we reveal that Argentine ants can discriminate between hydrocarbons with different branching patterns and the same chain length, but not always between hydrocarbons with the same branching patterns but different chain length. Our data thus show that biochemical characteristics influence those compounds that ants can discriminate between, and which thus potentially play a role in chemical signalling and nest-mate recognition.     

Cuticular hydrocarbons are heritable in the cricket Teleogryllus oceanicus

The ability of individuals to respond differentially to conspecifics depending on their genetic relatedness is a widespread phenomenon across the animal kingdom. Despite this, little is known about the selection processes that act on the phenotypic variation of traits used during recognition. Here we use a quantitative genetic approach to examine the patterns of genetic variation in cuticular hydrocarbon (CHC) profiles, a pheromonal system used extensively in insect communication. Using gas chromatography, we found family specificity in the CHC profiles of male crickets, Teleogryllus oceanicus. Across CHC peaks, our mean coefficient of additive genetic variation was 10.8%. Multivariate principal component analysis showed that most axes of variation were weighted by CHC peaks with significant additive genetic variation. Our results provide evidence that variation in CHC profiles can reflect genetic relatedness, supporting the widely held belief that this phenotypic trait is used as a mechanism for chemosensory kin recognition.     

Mating behavior and contact pheromones of Batocera horsfieldi (Hope) (Coleoptera: Cerambycidae)

The mating behavior of Batocera horsfieldi (Hope) was observed in the laboratory. The results showed that copulation of this longhorned beetle consisted of three phases: (i) encountering and pair‐bonding; (ii) mating attempt and ejaculation; and (iii) post‐copulatory guarding. Frozen females, with cuticular hydrocarbons stripped by hexane extraction, showed no attraction for males. Reapplying the solvent extract of frozen females to both washed dead males and females caused mating attempts by males, confirming that cuticular hydrocarbons (contact sex pheromones) played an important role in recognition of females by male B. horsfieldi.     

Behavioral and Chemical Correlates of Long-Term Queen Adoption in the Facultative Polygynous Ant Ectatomma tuberculatum

In ants, queen adoption is a common way of achieving secondary polygyny but the mechanisms involved are little known. Here we studied the process of long-term adoptions of alien queens in the facultative polygynous ant Ectatomma tuberculatum. In eight out of 10 successful adoption experiments, all the introduced queens showed similar behavior and fecundity as the resident queens even after 2 months, indicating complete integration into the colony. Chemical analysis revealed that the cuticular hydrocarbon profiles of resident and introduced queens were clearly distinct from those of workers and that they did not change after adoption. We propose that queen-specific cuticular hydrocarbon profile may represent a reliable signal of queen’s fertility and discuss about the evolution of high level of queen acceptance in E. tuberculatum.     

Development of interspecific recognition behavior in the antsManica rubida andFormica selysi (Hymenoptera: Formicidae) reared in mixed-species groups

The purpose of this study was to determine the role of early social experience on the ontogeny of kin and nestmate recognition in ants by means of both behavioral and chemical analysis. Workers of two ant species,Manica rubida (Myrmicinae) andFormica selysi (Formicinae), were reared in homospecific groups (control) or in artificial heterospecific groups (mixed), created less than 5 h after their emergence. Recognition was evaluated between unfamiliar individuals of different species reared in control and mixed groups for periods of 5, 12, 24, 48, and 72 h and 8, 15, 30, 60, or 90 days after emergence. Heterospecifically reared individuals of both species gradually became tolerant of allospecific individuals from control groups. Moreover, homospecifically reared individuals did not aggress allospecific individuals reared in mixed groups. During the course of familiarization between the species, there were modifications of the chemical recognition signals. In mixed groups, hydrocarbon profiles of both species acquire gradually some of the components characteristic of their heterospecific nestmates. These experiments showed that allospecific recognition required the acquisition of a minimal quantity of allospecific cues. The phenomenon provided an another example of the relationship between tolerance and the chemical cues displayed by both species. The results suggested that the individual recognized the allospecific cues borne on each individual's body surface and/or that each individual learned and memorized allospecific cues during its early life. Therefore, each individual might develop a template encoding the allospecific and the conspecific cues to characterize nestmates.     

A chemical level in the coevolutionary arms race between an ant social parasite and its hosts

Here we investigate the coevolutionary interactions between the slavemaking ant Protomognathus americanus and its Temnothorax hosts on a chemical level. We show that, although this social parasite is principally well-adapted to its hosts' cuticular hydrocarbon profile, there are pronounced differences in the fine-tuning of this adaptation. Between populations, chemical adaptation varies with host community composition, as the parasite faces a trade-off when confronted with more than one host species. In addition to adaptation of its own chemical signature, the slavemaker causes a reciprocal adjustment in its slaves' cuticular profile, the degree of which depends on the slave species. On the host side, successful parasite defence requires efficient enemy recognition, and in behavioural aggression trials, host colonies could indeed discriminate between invading slaves, which commonly accompany slavemakers on raids, and free-living conspecifics. Furthermore, hosts shifted their acceptance threshold over the seasons, presumably to reduce the costs of defence.     

Kin-informative recognition cues in ants

Although social groups are characterized by cooperation, they are also often the scene of conflict. In non-clonal systems, the reproductive interests of group members will differ and individuals may benefit by exploiting the cooperative efforts of other group members. However, such selfish behaviour is thought to be rare in one of the classic examples of cooperation--social insect colonies--because the colony-level costs of individual selfishness select against cues that would allow workers to recognize their closest relatives. In accord with this, previous studies of wasps and ants have found little or no kin information in recognition cues. Here, we test the hypothesis that social insects do not have kin-informative recognition cues by investigating the recognition cues and relatedness of workers from four colonies of the ant Acromyrmex octospinosus. Contrary to the theoretical prediction, we show that the cuticular hydrocarbons of ant workers in all four colonies are informative enough to allow full-sisters to be distinguished from half-sisters with a high accuracy. These results contradict the hypothesis of non-heritable recognition cues and suggest that there is more potential for within-colony conflicts in genetically diverse societies than previously thought.