Induced mimicry of colony odors in ants

The cuticular hydrocarbons ofFormica selysi (Formicinae) andMonica rubida (Myrmicinae) reared in single species and in mixed species colonies were determined using gas chromatography (GC) and GC-mass spectrometry. In colonies containing both species, each species modified its species-specific recognition odor. This odor is composed, at least partly, of cuticular hydrocarbons. The cuticular hydrocarbons ofM. rubida consist only of saturated alkanes (n-alkanes and branched alkanes). InF. selysi the mixture also contains unsaturated compounds (monoenes and dienes). In hetero-specific colonies, a new chemical signature developed. This signature resulted from qualitative and quantitative changes in the spectrum of hydrocarbons produced by each species and permitted the two species to inhabit the same nest without displaying interspecific aggression. The readjustment seemed to be more an active synthesis or an active transfer than simply a passive transfer from one species to the other. This may imply that the ants synthesized some components of the hydrocarbon signature of the other species. These synthesizing processes may be activated under particular social environmental conditions.     

Cuticular Hydrocarbon Composition Reflects Genetic Relationship Among Colonies of the Introduced Termite Reticulitermes santonensis Feytaud

Nestmate recognition plays a key role in kin selection to maintain colony integrity in social insects. Previous studies have demonstrated that nestmate recognition is dependent on detection of cuticular hydrocarbons. However, the absence of intraspecific aggression between some colonies of Isoptera and social Hymenoptera questions whether kin recognition must occur in social insects. The purpose of this study was to determine if cuticular hydrocarbon similarity and high genetic relatedness could explain the lack of intraspecific aggression among and within colonies of the introduced subterranean termite Reticulitermes santonensis. We performed both GC analysis of cuticular hydrocarbons and genotyping by using 10 DNA microsatellite loci on the same 10 workers from each of 14 parisian colonies. Multivariate analyses demonstrated correspondence between cuticular hydrocarbon patterns and genetic variation. By using a redundancy analysis combining chemical and genetic data, we found that a few hydrocarbons (mainly short vs. long chains; saturated vs. unsaturated alkanes) were associated with most genetic variation. We also found a strong positive correlation between chemical and genetic distances between colonies, thus providing evidence of a genetic basis for cuticular hydrocarbon variation. However, genetic distance did not account for all chemical variation, thus suggesting that some hydrocarbon variation was environmentally derived. Investigation at the intracolony level indicated that cuticular hydrocarbons did not depend on colony social structure. Based on our findings, we speculate that the absence of intraspecific aggression in R. santonensis may result from a loss of diversity in genetically derived recognition compounds in this species that presumably descended from R. flavipes populations imported from North America.     

Nestmate Recognition and the Role of Cuticular Hydrocarbons in the African Termite Raiding Ant <Emphasis Type="Italic">Pachycondyla analis</Emphasis>

Cuticular hydrocarbons (CHCs) are used for chemical communication among nestmates in many ant species, and they may play a role in the discrimination of nestmates and non-nestmates. Using the mandible opening response (MOR) bioassay, we tested the response of the African termite raiding ant, Pachycondyla analis, to CHC extracts of nestmates and non-nestmates. The ants were able to distinguish control chemical cues, from nestmate CHCs, and from non-nestmate CHCs, and, based on a CHC recognition threshold, aggression was demonstrated toward non-nestmates. Gas chromatography (GC) and GC-mass spectrometric analyses showed that CHC components of different ant colonies had chain lengths ranging from C₈ to C₃₁, comprising mainly n-alkanes, alkenes, and methyl branched alkanes, with the n-alkanes occurring in the same proportions among all colonies. The ants were grouped successfully according to their colonies of origin by using discriminant analysis of CHCs. We demonstrate that nestmate recognition occurs in P. analis, and that some of the cues involved are evidently alkenes and methyl-branched alkanes.     

Gaster flagging by fire ants (Solenopsis spp.): Functional significance of venom dispersal behavior

Behavioral and chemical studies with laboratory colonies indicate that the imported fire antSolenopsis invicta Buren (Myrmicinae) disperses venom through the air by raising and vibrating its gaster (i.e., gaster flagging). This mechanism of airborne venom dispersal is unreported for any ant species. Foraging workers utilize this air-dispersed venom (up to 500 ng) to repel heterospecifics encountered in the foraging arena, while brood tenders dispense smaller quantities ( 1 ng) to the brood surface, presumably as an antibiotic. Brood tenders removed from the brood cell and tested in heteropspecific encounters in the foraging arena exhibited the complete repertoire of agonistic gaster flagging behavior. These observations suggest that airborne venom dispersal by workers is context specific rather than temporal caste specific and that workers can control the quantity of venom released.     

Changes in the Cuticular Hydrocarbon Profile of the Slave-Maker Ant Queen, Polyergus breviceps Emery, After Killing a Formica Host Queen (Hymenoptera: Formicidae)

Queens of the slave-maker ant, Polyergus breviceps, take over nests of their Formica host species by fatally attacking the resident queen. As workers only begin grooming the P. breviceps queen once she has ceased her attack, we investigated whether a change in parasite queen chemistry may account for the change in worker behavior. Cuticular hydrocarbon profiles of newly mated P. breviceps queens and of queens of their two Formica host species were found to be species-specific. Profiles of newly mated P. breviceps queens that had attacked a Formica queen, however, were virtually identical to the queen profile of the species killed. Mass spectral analysis revealed that the hydrocarbons on the cuticles of newly mated P. breviceps changed from primarily normal alkanes to methyl and di-methyl branched alkanes after attacks. The results suggest that cuticular compounds from the host queen were transferred to the parasite queen during their aggressive interaction.     

Role of cuticular hydrocarbons of aphid parasitoids in their relationship to aphid-attending ants

Lysiphlebus cardui, the dominant aphidiid parasitoid of the black bean aphid,Aphis fabae cirsiiacanthoidis (Afc), on creeping thistle, is able to forage in ant-attended aphid colonies without being attacked by ants. Several behavioral observations and experimental studies led to the hypothesis thatL. cardui mimics the cuticular hydrocarbon profile of its host aphid. Chemical analysis of the cuticular extracts revealed that bothL. cardui and Afc exclusively possess saturated hydrocarbons:n-alkanes, monomethyl (MMA), dimethyl (DMA), and trimethyl alkanes (TMA). Comparison of the hydrocarbon profiles of parasitoid and aphid showed great qualitative resemblance between parasitoid and host:L. cardui possesses almost all host-specific compounds in addition to species-specific hydrocarbons of mainly higher molecular weight (>C₃₀). However, there is a lesser quantitative correspondence between parasitoid and host aphid. Furthermore, we analyzed the cuticular hydrocarbon profile of another parasitoid of Afc,Trioxys angelicae. This aphidiid species is vigorously attacked and finally killed by honeydewcollecting ants when encountered in aphid colonies. Its cuticular hydrocarbon profile is characterized by the presence of large amounts of (Z)-11-alkenes of chain lenghts C₂₇, C₂₉, C₃₁, and C₃₃, in addition to alkanes and presumably trienes. The role of the unsaturated hydrocarbons onT. angelicae as recognition cues for aphid-attending ants is discussed.     

Cuticular Hydrocarbons Provide Reliable Cues of Fertility in the Ant Gnamptogenys striatula

In ca. 150 species of queenless ants, a specialized queen caste is rare or absent, and mated workers take over the role of the queen in some or all of the colonies. Previously, it has been shown that reproduction in queenless ants is regulated by a combination of dominance behavior and chemical fertility signaling. It is unknown, however, whether chemical signals alone can sufficiently regulate reproduction. To investigate this possibility, we studied reproductive regulation in the facultatively queenless ant Gnamptogenys striatula, a species where dominance behavior is rare or absent. Active egg layers and infertile workers showed qualitative and quantitative differences in their cuticular hydrocarbon profile. Five long-chain methyl alkanes, 3,13- and 3,15-dimethyl pentriacontane, 3,13- and 3,15-dimethyl heptentriacontane, and 3,11,15-trimethyl heptentriacontane occurred only on the cuticles of virgin and mated egg layers. Pronounced quantitative differences were found in a further 27 alkenes; alkanes; and mono-, di-, and trimethyl alkanes. Workers that had recently stopped laying eggs had profiles similar to infertile workers, and mating status did not affect this chemical pattern. We conclude that the cuticular hydrocarbon profiles of G. striatula workers provide reliable information about their current fertility. In the interest of colony productivity, this allows reproduction to be regulated without the use of aggression.     

Inter- and Intraspecific Comparisons of Antiherbivore Defenses in Three Species of Rainforest Understory Shrubs

Plants defend themselves against herbivores and pathogens with a suite of morphological, phenological, biochemical, and biotic defenses, each of which is presumably costly. The best studied are allocation costs that involve trade-offs in investment of resources to defense versus other plant functions. Decreases in growth or reproductive effort are the costs most often associated with antiherbivore defenses, but trade-offs among different defenses may also occur within a single plant species. We examined trade-offs among defenses in closely related tropical rain forest shrubs (Piper cenocladum, P. imperiale, and P. melanocladum) that possess different combinations of three types of defense: ant mutualists, secondary compounds, and leaf toughness. We also examined the effectiveness of different defenses and suites of defenses against the most abundant generalist and specialist Piper herbivores. For all species examined, leaf toughness was the most effective defense, with the toughest species, P. melanocladum, receiving the lowest incidence of total herbivory, and the least tough species, P. imperiale, receiving the highest incidence. Although variation in toughness within each species was substantial, there were no intraspecific relationships between toughness and herbivory. In other Piper studies, chemical and biotic defenses had strong intraspecific negative correlations with herbivory. A wide variety of defensive mechanisms was quantified in the three Piper species studied, ranging from low concentrations of chemical defenses in P. imperiale to a complex suite of defenses in P. cenocladum that includes ant mutualists, secondary metabolites, and moderate toughness. Ecological costs were evident for the array of defensive mechanisms within these Piper species, and the differences in defensive strategies among species may represent evolutionary trade-offs between costly defenses.     

The Mandibular Gland Secretions of the Leaf-Cutting Ants Atta sexdens sexdens and Atta opaciceps Exhibit Intercaste and Intercolony Variations

The mandibular gland secretions of worker castes from wild colonies of the leaf-cutting ants Atta sexdens sexdens and Atta opaciceps were analyzed quantitatively by gas chromatography–mass spectrometry. The secretions comprised a complex mixture of volatile, mainly oxygenated compounds, and their profiles exhibited considerable qualitative and quantitative variations among species and castes. The known alarm pheromone 4-methyl-3-heptanone was common to both species. The elevated relative proportions of this ketone found in the secretions of gardeners and generalists suggest that such castes are primarily responsible for the production and release of the alarm pheromone. Quantitative variations (but no qualitative differences) in the profiles of secretions of soldiers from different colonies of A. sexdens sexdens were detected, supporting the view that intraspecific colony recognition is mediated through mandibular gland secretions. Subsequent laboratory assays showed that, among the compounds identified by GC–MS, 4-methyl-3-heptanone elicited a strong alarm response in workers of A. sexdens sexdens and A. opaciceps.     

Recognition of aphid parasitoids by honeydew-collecting ants: The role of cuticular lipids in a chemical mimicry system

The aphidiid waspLysiphlebus cardui parasitizes in ant-attendedAphis fabae cirsiiacanthoidis colonies without causing aggressive behavior in the antLasius niger. By contrast,Trioxys angelicae, another aphidiid parasitoid of aphids, is rapidly recognized and vigorously attacked by the ants.L. niger workers also responded differently to dead individuals ofL. cardui andT. angelicae. DeadL. cardui parasitoids were often ignored when encountered byL. niger, whereas deadT. angelicae individuals were immediately grasped by ants that discovered them. However, hexane-washed parasitoids caused a similar reaction pattern in the ants, in that both aphidiid species were tolerated in the aphid colony. Lure experiments demonstrated that chemical stimuli on the cuticle are major cues for the ants to distinguish between the parasitoids. The hexane extract ofL. cardui transferred to washed individuals ofT. angelicae resulted in ant responses characteristic towardsL. cardui, andL. niger workers displayed the typical removal pattern they normally showed towardsT. angelicae whenT. angelicae extract was applied toL. cardui individuals. Both parasitoid species treated with the hexane extract ofA. fabae cirsiiacanthoidis were similarily treated by the ants as were aphid control individuals. The suggestion that the aphidiid waspL. cardui uses chemical mimicry is discussed.     

Chemistry and functions of exocrine secretions of the antsTapinoma melanocephalum and T. erraticum

Volatile constituents produced by ant workers belonging to the speciesTapinoma melanocephalum andT. erraticum have been analyzed by gas chromatography-mass spectrometry. The pygidial (=anal) gland secretion ofT. melanocephalum is fortified with 6-methyl-5-hepten-2-one and actinidine (the mass spectrum of which is corrected in this paper). An unidentified compound was detected in cephalic extracts. The pygidial gland secretion ofT. erraticum was also dominated by 6-methyl-5-hepten-2-one, in addition to two isomers of iridodial, and iridomyrmecin. The sternal glands contained iridodial and C₁₅-C₂₀ hydrocarbons. Workers ofT. melanocephalum effectively utilize their pygidial gland secretions as an alarm-defense system during aggressive encounters with workers ofSolenopsis geminata. 6-Methyl-5-hepten-2-one is active as a releaser of alarm behavior, and actinidine is repellent to workers ofT. melanocephalum. Cephalic extracts possessed attractant and arrestant properties for workers of this species.     

Task-Related Differences in the Cuticular Hydrocarbon Composition of Harvester Ants, Pogonomyrmex barbatus

Colonies of the harvester ant, Pogonomyrmex barbatus, perform a variety of tasks. The behavior of an individual worker appears to depend on its recent history of brief contacts with ants of the same and other task groups. The purpose of this study was to determine whether task groups differ in cuticular hydrocarbon composition. We compared the cuticular hydrocarbon composition of ants collected under natural conditions as they performed one of three tasks: patrolling (locating food sources), foraging, or nest maintenance. Task groups differed significantly in the relative proportions of classes of hydrocarbon compounds, as well as in individual compounds. Relative to nest maintenance workers, foragers and patrollers had a higher proportion of straight-chain alkanes relative to monomethylalkanes, dimethylalkanes, and alkenes. There was no significant difference in the chain length of n-alkanes among the task groups. Foragers did not differ in hydrocarbon composition from patrollers. Colonies differed significantly from one another in hydrocarbon composition, but task groups differed in consistent ways from colony to colony, suggesting that the mechanism responsible for task-related hydrocarbon composition was the same in all colonies. P. barbatus workers switch tasks during their lifetimes, suggesting that cuticular hydrocarbon composition changes during adulthood as well. Nest maintenance workers are probably younger than foragers and patrollers and perform very little of their work outside of the nest. Task-related hydrocarbon differences detected here may be associated with worker age, and/or the abiotic characteristics (temperature, humidity, and ultraviolet light) of the interior and exterior work environments.     

Genetic Relatedness and Chemical Profiles in an Unusually Peaceful Eusocial Bee

Colonies of the stingless bee Tetragonilla collina frequently occur in unusually high densities and in direct neighborhood (nest aggregations), in rainforests of Southeast Asia. To investigate whether close relatedness and/or similar chemical profiles facilitate the co-occurrence of multiple T. collina colonies, we investigated aggressive behavior, genetic relatedness and cuticular hydrocarbon (CHC) profiles within and between colonies and nest aggregations. Although 17 out of 19 colonies within aggregations were largely unrelated, intraspecific aggression between different colonies was basically absent both within and among aggregations. This lack of aggression should favor social parasitism and hence the occurrence of unrelated individuals within a colony. However, low within-colony relatedness was found in only five out of 19 colonies where it may be explained by queen turnover or the occurrence of foreign workers. CHC profiles of colonies within and among aggregations were statistically different. However, many workers could chemically not be assigned to their maternal colony, indicating considerable overlap among colonies in odor profiles of workers. Moreover, odor profiles tended to be more similar within than among aggregations, although most colonies were unrelated. Thus, CHC profiles were a poor indicator of relatedness in T. collina. The lack of correlation between relatedness and chemical similarity in T. collina may be explained by the incorporation of resin-derived terpenes in their CHC profiles. The composition of these terpenes was highly similar among colonies, particularly within aggregations, hence potentially decreasing chemical distinctiveness and increasing behavioral tolerance.     

Changes in Monoterpene Emission Rates of <Emphasis Type="Italic">Quercus ilex</Emphasis> Infested by Aphids Tended by Native or Invasive <Emphasis Type="Italic">Lasius</Emphasis> Ant Species

The emission of volatile organic compounds (VOCs) depends on temperature and light. Other factors such as insect herbivory also may modify VOC emission. In particular, aphid feeding promotes the release of new compounds and changes the composition of plant volatile blends. Given that some aphids are tended by ants, we investigated whether ants change the emission of VOCs indirectly through attendance on aphids. The effect of Lachnus roboris aphids and two different tending ant species on terpene emission rates of 4-year-old holm oak (Quercus ilex) saplings was investigated during a field experiment. There were five treatments: saplings alone (T1), saplings infested with L. roboris aphids (T2), saplings infested with aphids tended by the local ant Lasius grandis (T3), those tended by small colonies of the invasive ant Lasius neglectus (T4), and those tended by large colonies of the same invasive ant species (T5). The infestation by L. roboris elicited the emission of Δ³-carene and increased the emission of myrcene and γ-terpinene. Terpene emissions were modified depending on the tending ant species. Attendance by the local ant L. grandis increased α and β-pinene and sabinene. Attendance by the invasive ant L. neglectus only decreased significantly the emission of myrcene, one of the major compounds of the Q. ilex blend. Aphid abundance decreased with time for all treatments, but there was no difference in aphid abundance among treatments. Total terpene emission rates were not correlated with aphid abundance. These results highlight that aphids and tending ants may change terpene emission rates, depending on the ant species.     

Dufour Gland Contents of Ants of the Cataglyphis bicolor Group

The species of desert-dwelling ants of the Cataglyphis bicolor (Hymenoptera: Formicidae) group are difficult to distinguish by morphological features. Analysis of the secretion from the Dufour glands of workers of a number of colonies was undertaken to see if it provided a clear test of species. Linked 6c-ms showed in all samples straight and branched-chain alkanes, linear alkenes, ketones, aldehydes, acetates, and a group of C₂₂ to C₂₈ esters not previously identified in this genus. Contents of the Dufour glands of C. savignyi from Tunisia and Egypt were similar, and comprised straight and branched-chain alkanes, alkenes and small amounts of esters. C. bicolor from Tunisia contained compounds similar to C. savignyi but was distinguished from the latter by larger amounts of the esters. The major compound in the glands of C. viaticus was tridecane, in contrast to the pentadecane of other species. It also contained a branched alkane, 3-methyltridecane as a major component. Branched-chain esters and a wide variety of acetates were also found in this species. C. diehlii had a limited range of compounds, with branched alkanes almost completely absent and high proportions of pentadecene and dodecyl acetate. C. bombycinus, a sympatric species, but recognized as not belonging to the bicolor group by its different mandibular gland substances, was notable in having butanoate esters in its Dufour glands. Despite these differences among species, both the great variability of individuals from a single colony and the among between conspecific colonies make species diagnosis from a few individuals difficult, in contrast with postpharyngeal glands, which, as recently reported, give a clearer indication of species.     

Novel internally branched,internal alkenes as major components of the cuticular hydrocarbons of the primitive australian antNothomyrmecia macrops Clark (Hymenoptera: Formicidae)

Internally mono- and dimethyl branched, internal alkenes, which constitute most of the cuticular hydrocarbon present on workers of the primitive Australian antNothomyrmecia macrops Clark, have been identified by gas chromatography-mass spectrometry. They are the first such alkenes reported from insects. Also present are alkanes with similar carbon skeletons. The hydrocarbon patterns of three separateNothomyrmecia colonies were very similar.     

Cuticular hydrocarbons of the paper wasp,Polistes fuscatus: A search for recognition pheromones

The cuticular chemicals of 124 individual wasps (foundresses and workers) from 23 colonies ofPolistes fuscatus were analyzed. The compounds identified, all of which were hydrocarbons, were similar to those of other vespid wasps in that the bulk of the hydrocarbons were 23–33 carbons in chain length. However, the hydrocarbon profile ofP. fuscatus differed from those of its congeners in its proportions of straight-chain alkanes, methylalkanes, and alkenes. Three of the 20 identified hydrocarbons, 13- and 15-MeC₃₁, 11,15- and 13,17-diMeC₃₁, and 13-, 15-, and 17-MeC₃₃, had properties postulated for recognition pheromones: colony specificity, efficacy in assigning wasps to the appropriate colony, heritability, lack of differences between foundresses and workers, and distinctive stereochemistry.     

A Review of Ant Cuticular Hydrocarbons

We compared the published cuticular hydrocarbon (CHC) profiles of 78 ant species across 5 subfamilies. Almost 1,000 CHCs have been described for these species, composing 187 distinct homologous series and ten hydrocarbon groups. In descending order of occurrence were: n-alkanes > monomethylalkanes > dimethylalkanes > alkenes > dienes>> trimethylalkanes>> methylalkenes > methylalkadienes > trienes > tetramethylalkanes. Odd chain lengths and positions of methyl or double bonds at odd carbon numbers were far more numerous than even chain-length compounds or bond positions. Although each species possess its own unique pattern of CHCs, we found no association between CHC profile and phylogeny. The production of the biosynthetically complex compounds (e.g., methyl branched dienes) by the most primitive living ant suggests that the basic genetic architecture required to produce the rich diversity of CHCs was already present prior to their adaptive radiation. Unlike the ubiquitous n-alkanes and monomethylalkanes, there is a huge diversity of species-specific dimethylalkanes that makes them likely candidates for species and nest-mate discrimination signals.     

Nestmate and Task Cues are Influenced and Encoded Differently within Ant Cuticular Hydrocarbon Profiles

Insect cuticular hydrocarbons (CHCs) are primarily antidesiccation agents, but they also play crucial roles in intra- and interspecific communication, especially among social Hymenoptera. The complex CHC profiles of social insects have often been compared among individuals, kin, nestmates, colonies, and species. In the ant Formica exsecta, only the (Z)-9-alkene part of the CHC profile encodes the nestmate signal. Here, we showed that the other major part of the CHC profile with n-alkane components is influenced strongly by the task a worker performs (foraging vs nonforaging). This part of the profile is independent of the nestmate signal. Therefore, the CHC profile of F. exsecta workers is composed of two independent parts: a colony-specific (Z)-9-alkene profile under genetic influence and an environmentally influenced task-related n-alkane profile. The dissociating of the CHC profile into two or more independent parts has implications for the analysis and interpretation of past and future CHC studies.     

Reaction of Mutualistic and Granivorous Ants to Ulex Elaiosome Chemicals

It has been proposed that chemicals on plant elaiosomes aid seed detection by seed-dispersing ants. We hypothesized that the chemical interaction between ants and elaiosomes is more intimate than a generic attraction, and that elaiosome chemicals will attract mutualistic but not granivorous ant species. We investigated this by using two gorse species, Ulex minor and U. europaeus, and two associated ant species from European heathlands, the mutualist Myrmica ruginodis and the granivore Tetramorium caespitum. Behavioral studies were conducted with laboratory nests and foraging arenas. Both ants will take Ulex seeds, but while M. ruginodis showed increased antennation toward ether extracts of elaiosome surface chemicals compared with controls, T. caespitum showed no response. Elaiosome extracts were separated into seven lipid fractions. M. ruginodis showed increased antennation only toward the diglyceride fractions of both Ulex species, whereas T. caespitum showed no consistent reaction. This indicates that M. ruginodis can detect the elaiosome by responding to its surface chemicals, but T. caespitum is unresponsive to these chemicals. Responses to surface chemicals could increase the rate of seed detection in the field, and so these results suggest that Ulex elaiosomes produce chemicals that facilitate attraction of mutualistic rather than granivorous ant species. This could reduce seed predation and increase Ulex fitness.