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.     

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.     

Acoustic alarm signalling facilitates predator protection of treehoppers by mutualist ant bodyguards

Mutualism is a net positive interaction that includes varying degrees of both costs and benefits. Because tension between the costs and benefits of mutualism can lead to evolutionary instability, identifying mechanisms that regulate investment between partners is critical to understanding the evolution and maintenance of mutualism. Recently, studies have highlighted the importance of interspecific signalling as one mechanism for regulating investment between mutualist partners. Here, we provide evidence for interspecific alarm signalling in an insect protection mutualism and we demonstrate a functional link between this acoustic signalling and efficacy of protection. The treehopper Publilia concava Say (Hemiptera: Membracidae) is an insect that provides ants with a carbohydrate-rich excretion called honeydew in return for protection from predators. Adults of this species produce distinct vibrational signals in the context of predator encounters. In laboratory trials, putative alarm signal production significantly increased following initial contact with ladybeetle predators (primarily Harmonia axyridis Pallas, Coleoptera: Coccinellidae), but not following initial contact with ants. In field trials, playback of a recorded treehopper alarm signal resulted in a significant increase in both ant activity and the probability of ladybeetle discovery by ants relative to both silence and treehopper courtship signal controls. Our results show that P. concava treehoppers produce alarm signals in response to predator threat and that this signalling can increase effectiveness of predator protection by ants.     

Evolution of cuticular hydrocarbon diversity in ants

The cuticular hydrocarbons (CHCs) of ants provide important cues for nest-mate and caste recognition. There is enormous diversity in the composition of these CHCs, but the manner in which this diversity has evolved is poorly understood. We gathered data on CHC profiles for 56 ant species, relating this information to their phylogeny. We deduced the mode of evolution of CHC profiles by reconstructing character evolution and then relating the number of changes in CHC components along each branch of the phylogeny to the length of the branch. There was a strong correlation between branch length and number of component changes, with fewer changes occurring on short branches. Our analysis thereby indicated a gradual mode of evolution. Different ant species tend to use specific CHC structural types that are exclusive of other structural types, indicating that species differences may be generated in part by switching particular biosynthetic pathways on or off in different lineages. We found limited, and contradictory, evidence for abiotic factors (temperature and rainfall) driving change in CHC profiles.     

Beauty or function? The opposing effects of natural and sexual selection on cuticular hydrocarbons in male black field crickets

Although many theoretical models of male sexual trait evolution assume that sexual selection is countered by natural selection, direct empirical tests of this assumption are relatively uncommon. Cuticular hydrocarbons (CHCs) are known to play an important role not only in restricting evaporative water loss but also in sexual signalling in most terrestrial arthropods. Insects adjusting their CHC layer for optimal desiccation resistance is often thought to come at the expense of successful sexual attraction, suggesting that natural and sexual selection are in opposition for this trait. In this study, we sampled the CHCs of male black field crickets (Teleogryllus commodus) using solid-phase microextraction and then either measured their evaporative water loss or mating success. We then used multivariate selection analysis to quantify the strength and form of natural and sexual selection targeting male CHCs. Both natural and sexual selection imposed significant linear and stabilizing selection on male CHCs, although for very different combinations. Natural selection largely favoured an increase in the total abundance of CHCs, especially those with a longer chain length. In contrast, mating success peaked at a lower total abundance of CHCs and declined as CHC abundance increased. However, mating success did improve with an increase in a number of specific CHC components that also increased evaporative water loss. Importantly, this resulted in the combination of male CHCs favoured by natural selection and sexual selection being strongly opposing. Our findings suggest that the balance between natural and sexual selection is likely to play an important role in the evolution of male CHCs in T. commodus and may help explain why CHCs are so divergent across populations and species.     

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.     

Changes of species specific cuticular hydrocarbon profiles in the cockroaches Nauphoeta cinerea and Leucophaea maderae reared in heterospecific groups

The cuticular hydrocarbon profiles of the cockroaches Nauphoeta cinerea and Leucophaea maderae are species-specific when maintained in homospecific rearings. When individuals were reared in mixed species colonies, they initially remained in homospecific groups under different shelters. However, after 14 days they formed one heterospecific group with cuticular profiles showing characteristics of both species. When individuals were returned in monospecific rearings, their cuticular hydrocarbon profiles returned to species-specific ones within 3 weeks.     

Cuticular hydrocarbons influence female attractiveness to males in the Australian field cricket,Teleogryllus oceanicus

Sexual dimorphism is thought to result from directional sexual selection acting on male signal traits, with female signal traits given little, if any, attention. Here, we examine male mating preferences in the Australian field cricket, Teleogryllus oceanicus. Using a multivariate selection analysis approach, we found that male preferences have the potential to exert selection on female cuticular hydrocarbons, chemical compounds widely used as sexual signals in insects. In addition to finding both stabilizing and disruptive preference gradients, we also found weak negative directional preference for female cuticular hydrocarbons. We contrast our results with a recent study examining sexual selection via female choice on male T. oceanicus cuticular hydrocarbons and suggest that differences in the form and intensity of sexual selection between the genders may provide part of the net selection differential necessary for the evolution of sexual dimorphism in this species.     

Striking cuticular hydrocarbon dimorphism in the mason wasp Odynerus spinipes and its possible evolutionary cause (Hymenoptera: Chrysididae,Vespidae)

Cleptoparasitic wasps and bees smuggle their eggs into the nest of a host organism. Here the larvae of the cleptoparasite feed upon the food provision intended for the offspring of the host. As cleptoparasitism incurs a loss of fitness for the host organism (offspring of the host fail to develop), hosts of cleptoparasites are expected to exploit cues that alert them to potential cleptoparasite infestation. Cuticular hydrocarbons (CHCs) could serve as such cues, as insects inevitably leave traces of them behind when entering a nest. By mimicking the host''s CHC profile, cleptoparasites can conceal their presence and evade detection by their host. Previous studies have provided evidence of cleptoparasites mimicking their host''s CHC profile. However, the impact of this strategy on the evolution of the host''s CHC profile has remained unexplored. Here, we present results from our investigation of a host–cleptoparasite system consisting of a single mason wasp species that serves syntopically as the host to three cuckoo wasp species. We found that the spiny mason wasp (Odynerus spinipes) is able to express two substantially different CHC profiles, each of which is seemingly mimicked by a cleptoparasitic cuckoo wasp (i.e. Chrysis mediata and Pseudospinolia neglecta). The CHC profile of the third cuckoo wasp (Chrysis viridula), a species not expected to benefit from mimicking its host''s CHC profile because of its particular oviposition strategy, differs from the two CHC profiles of its host. Our results corroborate the idea that the similarity of the CHC profiles between cleptoparasitic cuckoo wasps and their hosts are the result of chemical mimicry. They further suggest that cleptoparasites may represent a hitherto unappreciated force that drives the evolution of their hosts'' CHCs.     

Agonistic relationships among sympatric mediterranean ant species (Hymenoptera: Formicidae)

The potential aggressiveness of nine ant species belonging to the same community was estimated by means of two laboratory approaches: individual confrontations and group confrontations. Interspecific aggressiveness was not directly related to size of species in individual tests, although in group tests mortality of smaller ants increased with increasing size difference between the opponents. Tempo was related to the defense strategy of species (low-tempo species used immobility as a mechanism to avoid attacks, while high-tempo species tended to escape when coming into contact with more aggressive ones), but not with their potential aggressiveness: the most aggressive ants in individual and group tests were both low-tempo and high-tempo species when only these two categories were considered. No specific level of aggression was related to the subfamily to which the species belong (both myrmicines and formicines dominated in individual and group tests) or to the degree of taxonomic similarity between species.     

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.