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Cuticular hydrocarbons (CHCs) constitute an important class of chemical compounds present in the cuticular layer of insects, where their main functions are to prevent desiccation of the cuticle and as signals for intraspecific recognition. Studies concerning CHCs have shown a phenotypic flexibility of their composition, depending on environmental factors. However, the way that each of these factors influences this variation remains little explored. The aim of the present study was to evaluate the effects of environmental variations on the cuticular chemical composition of the ant Atta laevigata. Workers from four different colonies nesting in forest edge environments were collected over the course of a year, during the hot and humid and cold and dry seasons. The cuticular compounds were extracted and then analyzed by gas chromatography, revealing that the compounds of this species belonged to the classes of linear alkanes, mono, di and trimethyl alkanes, alkenes and alkadienes. Furthermore, the cuticular profile varied significantly among colonies of this species and between seasons, while intra-season variability was more significant during the hot and humid season. The observed temporal variation indicated that the numbers of compounds and the proportion of different types of CHC can vary according to the period of the year, however with a greater significant variation in colonies in the hot and humid season. These results showed that variations in environmental conditions, especially climate, can have decisive effects in the dynamics of cuticular chemistry., Competing Interests: Declarations. Competing Interests: The authors declare no competing interests., (© 2025. The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature.)

Ants use chemical cues known as cuticular hydrocarbons (CHCs) for both intraspecific and interspecific recognition. These compounds serve ants in distinguishing between nestmates and non-nestmates, enabling them to coexist in polydomous colonies characterized by socially connected yet spatially separated nests. Hence, the aim of this study was to investigate the intraspecific aggression level between nestmates and non-nestmates of the bullet ant Paraponera clavata (Fabricius, 1775), analyze and compare their CHCs, and evaluate the occurrence of polydomy in this species. We conducted aggression tests between foragers, both in laboratory and field settings. To identify the chemical profiles, we utilized gas chromatography coupled with mass spectrometry (GC-MS). We marked the foragers found at nest entrances and subsequently recaptured these marked ants to validate workers exchange among nests. Across all nests, a low intraspecific aggression level was observed within the same area. However, a significant difference in aggression correlated to distance between nests. Analysis of the cuticular chemical profile of P. clavata unveiled colony-specific CHCs, both qualitatively and quantitatively. Notably, we observed instances of ants from certain nests entering or exiting different nests. This behavior, in conjunction with the observed low intraspecific aggression despite differences in CHCs suggests polydomy for this species. Polydomy can offer several benefits, including risk spreading, efficient exploitation of resources, potential for colony size increasing and reduced costs associated with foraging and competition., (© 2024. The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature.)

Nestmate recognition in ants is based on perceived differences in a multi-component blend of hydrocarbons that are present on the insect cuticle. Although supplementation experiments have shown that some classes of hydrocarbons, such as methyl branched alkanes and alkenes, have a salient role in nestmate recognition, there was basically no information available on how ants detect and perceive these molecules. We used a new conditioning procedure to investigate whether individual carpenter ants could associate a given hydrocarbon (linear or methyl-branched alkane) to sugar reward. We then studied perceptual similarity between a hydrocarbon previously associated with sugar and a novel hydrocarbon. Ants learnt all hydrocarbon-reward associations rapidly and with the same efficiency, regardless of the structure of the molecules. Ants could discriminate among a large number of pairs of hydrocarbons, but also generalised. Generalisation depended both on the structure of the molecule and the animal's experience. For linear alkanes, generalisation was observed when the novel molecule was smaller than the conditioned one. Generalisation between pairs of methyl-alkanes was high, while generalisation between hydrocarbons that differed in the presence or absence of a methyl group was low, suggesting that chain length and functional group might be coded independently by the ant olfactory system. Understanding variations in perception of recognition cues in ants is necessary for the general understanding of the mechanisms involved in social recognition processes based on chemical cues., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Social insects offer unique opportunities to test predictions regarding the evolution of cooperation, life histories and communication. Colony founding by groups of unrelated queens, some of which are later killed, may select for selfish reproductive strategies, honest signalling and punishment. Here, we use a brood transfer experiment to test whether cofounding queens of the ant Lasius niger 'selfishly' adjust their productivity when sharing the nest with future competitors. We simultaneously analysed queen cuticular hydrocarbon (CHC) profiles to investigate whether queens honestly signal their reproductive output or produce dishonest, manipulative signals, providing a novel test of the evolutionary significance of queen pheromones. Queens produced fewer workers when their colony contained ample brood, but only in the presence of competitors, suggesting selfish conservation of resources. Several CHCs correlated with reproductive maturation, and to a lesser extent with productivity; the same hydrocarbons were more abundant on queens that were not killed, suggesting that workers select productive queens using these chemical cues. Our results highlight the role of honest signalling in the evolution of cooperation: whenever cheaters can be reliably identified, they may incur sanctions that reduce the incentive to be selfish.

Cuticular hydrocarbon (CHC) profiling shows promise as a chemotaxonomic tool for identifying and discriminating between closely related insect species. However, there have been limited studies using CHC profiling to differentiate between weevil species (Coleoptera: Curculionidae). This proof-of-concept study investigated the use of CHC and volatile profiling to discriminate between five weevil species from three genera in the Gonipterini tribe. A total of 56 CHCs and 41 other volatile compounds were found across the five species, with 83 of the compounds being identified through their mass fragmentation patterns. The number of CHCs from each species ranged from 20 to 43, while the proportion of CHCs unique to each species varied between 0% and 19%. The most abundant CHCs were nonacosane, 7-methylheptacosane, heptacosane, and hexacosane. Principal component analysis of the centred log-ratio transformed data revealed broad differences in CHC profiles between the two Oxyops species, with Bryachus squamicollis demonstrating the greatest divergence from the other Gonipterini species. The results suggest that CHC analysis could be used to support established taxonomic methods, including morphological features and genetic sequencing results. [ABSTRACT FROM AUTHOR]

Organismal traits may experience conflicting selection pressures if they fulfil different functions simultaneously. This can require trade‐offs between functions or alternatively functional separation between elements of the trait.An important multifunctional trait in insects is the cuticular hydrocarbon (CHC) layer. CHCs cover the body of nearly all insects, protect against desiccation and serve as a communication signal. In social insects like ants, they provide cues for nestmate recognition. To maintain their waterproofing function, insects have to adjust CHC composition to current temperatures. These changes might affect information content and interfere with communication, which would be especially detrimental in social insects.Here, we studied how acclimation affects nestmate recognition in two sister species of the ant genus Lasius. Colony fragments were exposed to three climate regimes. We analysed behaviour towards same and differently acclimated conspecifics, and determined which CHCs were related to acclimatory changes, colony differences and inter‐individual aggression.Differential acclimation led to higher aggression and chemical distances among former nestmates. We identified small CHC subsets, which only differed among colonies or among acclimation treatments. Moreover, few compounds sufficed to explain inter‐individual aggression, suggesting that ants do not use the entire CHC profile for nestmate recognition and that colony identity is encoded in a redundant way.Across individual CHCs, their contribution to colony differences and to differences among acclimatory treatments was negatively correlated, indicating that there is some degree of functional separation. However, CHC classes could not be clearly assigned to one or another function, indicating that the role of each CHC is idiosyncratic and may differ among species. Acclimatory effects and colony differences were more independent from each other in L. platythorax than in L. niger, indicating that functional separation can differ even among sister species.Our results show that CHC functions are more intertwined than previously assumed, suggesting that insects cannot optimise all functions independently. The main constraint might be the need to maintain a certain phase behaviour of the CHC layer, which depends on CHC composition and affects functionality. The need to separate functions might depend on species‐specific ecological and life‐history parameters.Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

A series of novel pleuromutilin derivatives were designed and synthesized based on the twin drugs theory. Piperazinyl and thioether were used as the linkage to connect the pleuromutilin nuclear and sulfonamide to improve the biological activity and water solubility of derivatives. The in vitro antibacterial activities against drug-sensitive and drug-resistance bacteria were evaluated by agar dilution method. Most of the 25 compounds displayed excellent antibacterial activities against drug-sensitive bacteria, particularly, five compounds (9, 10, 11, 14a and 14b) exerted the excellent antibacterial activities against drug-resistance bacteria. [ABSTRACT FROM AUTHOR]

The production of royal pheromones by reproductives (queens and kings) enables social insect colonies to allocate individuals into reproductive and non-reproductive roles. In many termite species, nestmates can develop into neotenics when the primary king or queen dies, which then inhibit the production of additional reproductives. This suggests that primary reproductives and neotenics produce royal pheromones. The cuticular hydrocarbon heneicosane was identified as a royal pheromone in Reticulitermes flavipes neotenics. Here, we investigated the presence of this and other cuticular hydrocarbons in primary reproductives and neotenics of this species, and the ontogeny of their production in primary reproductives. Our results revealed that heneicosane was produced by most neotenics, raising the question of whether reproductive status may trigger its production. Neotenics produced six additional cuticular hydrocarbons absent from workers and nymphs. Remarkably, heneicosane and four of these compounds were absent in primary reproductives, and the other two compounds were present in lower quantities. Neotenics therefore have a distinct 'royal' blend from primary reproductives, and potentially over-signal their reproductive status. Our results suggest that primary reproductives and neotenics may face different social pressures. Future studies of these pressures should provide a more complete understanding of the mechanisms underlying social regulation in termites. [ABSTRACT FROM AUTHOR]

In social insects, cuticular hydrocarbons function in nest-mate recognition and also provide a waxy barrier against desiccation, but basic evolutionary features, including the heritability of hydrocarbon profiles and how they are shaped by natural selection are largely unknown. We used a new pharaoh ant (Monomorium pharaonis) laboratory mapping population to estimate the heritability of individual cuticular hydrocarbons, genetic correlations between hydrocarbons, and fitness consequences of phenotypic variation in the hydrocarbons. Individual hydrocarbons had low to moderate estimated heritability, indicating that some compounds provide more information about genetic relatedness and can also better respond to natural selection. Strong genetic correlations between compounds are likely to constrain independent evolutionary trajectories, which is expected, given that many hydrocarbons share biosynthetic pathways. Variation in cuticular hydrocarbons was associated with variation in colony productivity, with some hydrocarbons experiencing strong directional selection. Altogether, this study builds on our knowledge of the genetic architecture of the social insect hydrocarbon profile and indicates that hydrocarbon variation is shaped by natural selection. [ABSTRACT FROM AUTHOR]

Colony coherence is essential for eusocial insects because it supports the inclusive fitness of colony members. Ants quickly and reliably recognize who belongs to the colony (nestmates) and who is an outsider (non-nestmates) based on chemical recognition cues (cuticular hydrocarbons: CHCs) which as a whole constitute a chemical label. The process of nestmate recognition often is described as matching a neural template with the label. In this study, we tested the prevailing view that ants use commonalities in the colony odor that are present in the CHC profile of all individuals of a colony or whether different CHC profiles are learned independently. We created and manipulated sub-colonies by adding one or two different hydrocarbons that were not present in the original colony odor of our Camponotus floridanus colony and later tested workers of the sub-colonies in one-on-one encounters for aggressive responses. We found that workers adjust their nestmate recognition by learning novel, manipulated CHC profiles, but still accept workers with the previous CHC profile. Workers from a sub-colony with two additional components showed aggression against workers with only one of the two components added to their CHC profile. Thus, additional components as well as the lack of a component can alter a label as “non-nestmate.” Our results suggest that ants have multiple-templates to recognize nestmates carrying distinct labels. This finding is in contrast to what previously has been proposed, i.e., a widening of the acceptance range of one template. We conclude that nestmate recognition in ants is a partitioned (multiple-template) process of the olfactory system that allows discrimination and categorization of nestmates by differences in their CHC profiles. Our findings have strong implications for our understanding of the underlying mechanisms of colony coherence and task allocation because they illustrate the importance of individual experience and task associated differences in the CHC profiles that can be instructive for the organization of insect societies. [ABSTRACT FROM AUTHOR]

In insect societies, chemical communication plays an important role in colony reproduction and individual social status. Many studies have indicated that cuticular hydrocarbons (CHCs) are the main chemical compounds encoding reproductive status. However, these studies have largely focused on queenless or monogynous species whose workers are capable of egg laying and have mainly explored the mechanisms underlying queen-worker or worker-worker reproductive conflicts. Less is known about what occurs in highly polygynous ant species with permanently sterile workers. Here, we used the Argentine ant as a model to examine the role of CHCs in communicating reproductive information in such insect societies. The Argentine ant is unicolonial, highly polygynous, and polydomous. We identified several CHCs whose presence and levels were correlated with queen age, reproductive status, and fertility. Our results also provide new insights into queen executions in the Argentine ant, a distinctive feature displayed by this species in its introduced range. Each spring, just before new sexuals appear, workers eliminate up to 90% of the mated queens in their colonies. We discovered that queens that survived execution had different CHC profiles from queens present before and during execution. More specifically, levels of some CHCs were higher in the survivors, suggesting that workers could eliminate queens based on their chemical profiles. In addition, queen CHC profiles differed based on season and species range (native vs. introduced). Overall, the results of this study provide new evidence that CHCs serve as queen signals and do more than just regulate worker reproduction. [ABSTRACT FROM AUTHOR]

Social insects recognize their nestmates by means of a cuticular hydrocarbon signature shared by colony members, but how nest signature changes across time has been rarely tested in longitudinal studies and in the field. In social wasps, the chemical signature is also deposited on the nest surface, where it is used by newly emerged wasps as a reference to learn their colony odor. Here, we investigate the temporal variations of the chemical signature that wasps have deposited on their nests. We followed the fate of the colonies of the social paper wasp Polistes biglumis in their natural environment from colony foundation to decline. Because some colonies were invaded by the social parasite Polistes atrimandibularis, we also tested the effects of social parasites on the nest signature. We observed that, as the season progresses, the nest signature changed; the overall abundance of hydrocarbons as well as the proportion of longer-chain and branched hydrocarbons increased. Where present, social parasites altered the host-nest signature qualitatively (adding parasite-specific alkenes) and quantitatively (by interfering with the increase in overall hydrocarbon abundance). Our results show that 1) colony odor is highly dynamic both in colonies controlled by legitimate foundresses and in those controlled by social parasites; 2) emerged offspring contribute little to colony signature, if at all, in comparison to foundresses; and 3) social parasites, that later mimic host signature, initially mark host nests with species-specific hydrocarbons. This study implies that important updating of the neural template used in nestmate recognition should occur in social insects. [ABSTRACT FROM AUTHOR]

Cuticular hydrocarbons (CHCs) cover the cuticles of virtually all insects, serving as a waterproofing agent and as a communication signal. The causes for the high CHC variation between species, and the factors influencing CHC profiles, are scarcely understood. Here, we compare CHC profiles of ant species from seven biogeographic regions, searching for physiological constraints and for climatic and biotic selection pressures. Molecule length constrained CHC composition: long-chain profiles contained fewer linear alkanes, but more hydrocarbons with disruptive features in the molecule. This is probably owing to selection on the physiology to build a semi-fluid cuticular layer, which is necessary for waterproofing and communication. CHC composition also depended on the precipitation in the ants' habitats. Species from wet climates had more alkenes and fewer dimethyl alkanes than those from drier habitats, which can be explained by different waterproofing capacities of these compounds. By contrast, temperature did not affect CHC composition. Mutualistically associated (parabiotic) species possessed profiles highly distinct from non-associated species. Our study is, to our knowledge, the first to show systematic impacts of physiological, climatic and biotic factors on quantitative CHC composition across a global, multi-species dataset. We demonstrate how they jointly shape CHC profiles, and advance our understanding of the evolution of this complex functional trait in insects. [ABSTRACT FROM AUTHOR]

Highly social ants, bees and wasps employ sophisticated recognition systems to identify colony members and deny foreign individuals access to their nest. For ants, cuticular hydrocarbons serve as the labels used to ascertain nest membership. Social parasites, however, are capable of breaking the recognition code so that they can thrive unopposed within the colonies of their hosts. Here we examine the influence of the socially parasitic slave-making ant, Polyergus breviceps on the nestmate recognition system of its slaves, Formica altipetens. We compared the chemical, genetic, and behavioral characteristics of colonies of enslaved and free-living F. altipetens. We found that enslaved Formica colonies were more genetically and chemically diverse than their free-living counterparts. These differences are likely caused by the hallmark of slave-making ant ecology: seasonal raids in which pupa are stolen from several adjacent host colonies. The different social environments of enslaved and free-living Formica appear to affect their recognition behaviors: enslaved Formica workers were less aggressive towards non-nestmates than were free-living Formica. Our findings indicate that parasitism by P. breviceps dramatically alters both the chemical and genetic context in which their kidnapped hosts develop, leading to changes in how they recognize nestmates. [ABSTRACT FROM AUTHOR]

Queen pheromones, which signal the presence of a fertile queen and induce daughter workers to remain sterile, are considered to play a key role in regulating the reproductive division of labor of insect societies. Although queen pheromones were long thought to be highly taxon-specific, recent studies have shown that structurally related long-chain hydrocarbons act as conserved queen signals across several independently evolved lineages of social insects. These results imply that social insect queen pheromones are very ancient and likely derived from an ancestral signalling system that was already present in their common solitary ancestors. Based on these new insights, we here review the literature and speculate on what signal precursors social insect queen pheromones may have evolved from. Furthermore, we provide compelling evidence that these pheromones should best be seen as honest signals of fertility as opposed to suppressive agents that chemically sterilize the workers against their own best interests. [ABSTRACT FROM AUTHOR]

Macaranga myrmecophytes (ant-plants) are generally well protected from herbivore attacks by their symbiotic ants (plant-ants). However, larvae of Arhopala (Lepidoptera: Lycaenidae) species survive and develop on specific Macaranga ant-plant species without being attacked by the plant-ants of their host species. We hypothesized that Arhopala larvae chemically mimic or camouflage themselves with the ants on their host plant so that the larvae are accepted by the plant-ant species of their host. Chemical analyses of cuticular hydrocarbons showed that chemical congruency varied among Arhopala species; A. dajagaka matched well the host plant-ants, A. amphimuta did not match, and unexpectedly, A. zylda lacked hydrocarbons. Behaviorally, the larvae and dummies coated with cuticular chemicals of A. dajagaka were well attended by the plant-ants, especially by those of the host. A. amphimuta was often attacked by all plant-ants except for the host plant-ants toward the larvae, and those of A. zylda were ignored by all plant-ants. Our results suggested that conspicuous variations exist in the chemical strategies used by the myrmecophilous butterflies that allow them to avoid ant attack and be accepted by the plant-ant colonies. [ABSTRACT FROM AUTHOR]

Caterpillars of the parasitic lycaenid butterfly are often adopted by host ants. It has been proposed that this adoption occurs because the caterpillars mimic the cuticular hydrocarbons of the host ant. This study aimed to examine whether caterpillars of the Japanese lycaenid butterfly Niphanda fusca induce adoption by mimicking their host ant Camponotus japonicus. Behavioral observations conducted in the laboratory showed that most second-instar caterpillars were not adopted, whereas most third-instar caterpillars were successfully adopted by host workers. A chemical comparison detected no characteristic differences in the cuticular hydrocarbon profiles between second- and third-instar caterpillars. However, morphological features of the caterpillars differed between the second and third instars; third-instar caterpillars developed exocrine glands (ant organs) such as tentacle organs and a dorsal nectary organ. These results suggest that multiple chemical signatures, not only cuticular hydrocarbons, may be important for invasion of the host ant nest. [ABSTRACT FROM AUTHOR]

The exoskeletons of insects are covered by complex mixtures of cuticular hydrocarbons (CHCs) which are involved in social and sexual communication. However, little is known about the relationship between the structures of CHCs and their behavioral activity. The key component of the contact sex pheromone of the parasitoid Lariophagus distinguendus is 3-methylheptacosane (3-MeC27), which is present in CHC profiles of both females and newly emerged males. The CHCs of females and young males elicit wing-fanning behavior in older males. However, as young males age, 3-MeC27 disappears from their CHC profiles and they no longer elicit wing-fanning responses from other males. We applied enantiopure 3-MeC27 and structurally related CHCs (with respect to chain length or methyl-branch position) to the cuticle of aged male dummies and recorded the wing-fanning behavior of responding males. Only the two enantiomers of 3-MeC27 restored the dummies' attractiveness. The addition of structurally related CHCs or various n-alkanes to bioactive dummies of young males and females significantly decreased wing-fanning by test males. Hence, L. distinguendus males respond specifically but not enantioselectively to 3-MeC27, and perceive the CHC profiles as a whole. Both removal (as is the case with 3-MeC27 in aging males) and addition of individual compounds may disrupt the behavioral response. [ABSTRACT FROM AUTHOR]

Communication between organisms involves visual, auditory, and olfactory pathways. In solitary insects, chemical recognition cues are influenced mainly by selection regimes related to species recognition and sexual selection. In social insects, chemical recognition cues have an additional role in mediating recognition of society members and, thereby, allowing kin selection to operate. Here, we examined whether cuticular hydrocarbon profiles are sex-specific and whether males and young queens of the ant Formica fusca have colony-specific profiles. We also investigated whether there is a relationship between genetic relatedness and chemical diversity within colonies. We demonstrated that female and male sexuals do not have unique sex-specific compounds, but that there are quantitative chemical differences between the sexes. Out of the 51 cuticular hydrocarbon compounds identified, 10 showed a significant quantitative difference between males and females. We also showed that both males and females have a significant colony-specific component in their profiles. Finally, we found a negative correlation between within-colony relatedness and within-colony chemical diversity of branched, but not linear compounds. This suggests that colonies with multiple matri- or patrilines also have a significantly greater chemical diversity. [ABSTRACT FROM AUTHOR]

In insects, cuticular hydrocarbons (CHCs) generally are used as cues and signals for within colony processes, such as signaling reproductive status, and between colony processes, such as colony membership. We examined CHC profiles of the facultatively polygynous ant Pachycondyla verenae in order to identify chemical signals of reproductive queens within colonies containing many gynes. Colonies of P. verenae, belonging to two different members of a complex of morphospecies, were collected from three geographic localities within South America. We also tested whether CHC profiles differed between geographic localities and morphospecies. We found three alkenes, two isomers of pentacosene and heptacosene, which were more abundant in CHC profiles of reproductive queens of this morphospecies complex. When we tested whether these differences were consistent across geographic localities, we found the abundance of these alkenes differed according to morphospecies, with the isomers of pentacosene being more abundant in queens from morph one, and heptacosene being more abundant in queens from morph two. Our study has given further insight into the mechanisms behind maintenance of reproductive dominance, and has demonstrated that chemical signatures associated with reproductive status in Pachycondyla verenae are not conserved within this species complex. [ABSTRACT FROM AUTHOR]

Cuticular hydrocarbons (CHCs) are expressed on an insect's cuticle and are one of the major factors allowing insects to identify members of their own species, colony and gender. As a result of their species-specificity, CHCs are increasingly used to delimit species in addition to more conventional methods, such as morphology or genetic markers, and so play an important role in chemotaxonomy. Species vary in the type of CHCs that they produce, as well as in the relative quantities of shared compounds. This review summarizes not only how taxonomists may differentiate between species based on CHC profiles, but also the incentive for using CHC composition as taxonomic tool. Benefits regarding the identification of cryptic species and early signs of reproductive isolation are then discussed, giving examples from studies of taxonomy, behaviour and biosynthesis. For taxonomic characters to reliably indicate species boundaries, their limitations need to be known. Potential problems caused by environmental effects, intra-species variation in profiles and other technical issues are highlighted, and suggestions are made regarding their avoidance. It remains a challenge to determine the variation beyond which two species can be called independent; a problem shared by most methods of delimitation. Recently, there has been a shift towards using a combination of different taxonomic tools, both molecular and non-molecular, to test observed species differences. [ABSTRACT FROM AUTHOR]

Summary 1. Trade-offs between current and future resource allocation can select for elevated reproductive effort in individuals facing mortality. Males are predicted to benefit from increasing investment in costly sexually selected signals after experiencing an acute life span reduction, although few examples of such facultative terminal investment are known. 2. In the mealworm beetle, Tenebrio molitor, males' odours become more attractive to females following a life-threatening immune challenge. However, the pheromones involved are unknown, hindering further insight into the proximate mechanisms and ultimate consequences of terminal investment. 3. Using chemical and behavioural analyses, we show that the cuticular hydrocarbons (CHCs) of T. molitor are sexually dimorphic and are used by females to locate and select males. Moreover, both male CHCs and glandular pheromones were affected by experimental immune challenge in a fashion that made them more attractive to females. 4. The results suggest that males terminally invest in both short- and medium-range pheromones when they perceive reduced future survival. Moreover, the constitutive and inducible aspects of male and female CHC production are consistent with sex-specific selection on the signalling and defensive functions of CHCs. The implications of terminal investment for 'dishonest' signalling and the efficacy of sexual selection are discussed. [ABSTRACT FROM AUTHOR]

In most ants, bees, and wasps, the workers are capable of challenging the reproductive monopoly of the queen by laying unfertilized, male eggs. An important mechanism that can resolve this conflict is policing, whereby the queen or workers prevent successful worker reproduction by selectively eating worker-laid eggs or by attacking egg-laying workers. Egg policing by workers has been shown to occur in several social wasp species, but the information used by worker wasps to discriminate between queen-laid and worker-laid eggs has never been investigated. Our aim, therefore, was to investigate if hydrocarbons might be used in egg policing by workers in the common wasp, Vespula vulgaris, where worker policing previously has been shown to be effective. Our results show that 51 different hydrocarbons are present on the surface of newly-laid eggs, and that there are pronounced quantitative differences in the hydrocarbon profiles of queen-laid and worker-laid eggs, with longer-chained alkenes and methylated alkanes (C-C) in particular being more abundant on the surface of queen-laid eggs. We further show that the hydrocarbon profiles on the surface of queen-laid and worker-laid eggs resemble those found on the mother queen's and workers' cuticles. Interestingly, longer-chained methylated alkanes also were more abundant on the cuticle of both mother queens and reproductive workers, suggesting that these compounds are linked to fertility, as has also been found to be the case in several ant species. [ABSTRACT FROM AUTHOR]

Split sex ratio theory predicts that when kin structure varies among colonies of social insects, in order to maximize the inclusive fitness, colonies with relatively high sister-sister relatedness should specialize in producing reproductive females, whereas in those with relatively low sister-sister relatedness workers should bias their sex ratio towards males. However, in order to achieve this, workers need to be able to reliably assess the type of colony in which they live. The information on colony kin structure may be encoded in cuticular hydrocarbons (CHCs), assuming that genetic variability translates accurately into chemical variability. However, in genetically heterogeneous colonies, too accurate information may encourage the pursuit of individual interests through nepotistic behavior and reduce colony efficiency or cause social disruption. In this study, we estimated how well variability of CHC recognition cues reflects colony kin structure in the ant Formica exsecta. Our results show that CHC variability does not covary with kin structure or the overall genetic diversity of the colony, and that patrilines and matrilines can have distinct CHC profiles in some but not all colonies. However, within-colony relatedness remains the key determinant of colony sex ratios. Based on our results, CHC variability cannot serve as accurate information on within-colony relatedness, kin structure, or full-sib affiliation, nor do workers seem to use colony CHC variability as a proxy for sex-ratio adjustment. The use of this type of information thus could lead workers to make mistakes, and it remains unclear how colonies of Formica exsecta adjust offspring sex ratio to their optimal value. [ABSTRACT FROM AUTHOR]

Cuticular hydrocarbons (CHCs) are increasingly recognized as important to insects and are used for constructing taxonomies. However, multiple parameters affect the expression of CHCs besides a genetic component. We propose that selection may act differently on the expression of CHCs, depending on the evolutionary context. To explore the influence of selection, the CHCs of two closely related ant species, Lasius niger and Lasius platythorax, were studied in a multidisciplinary approach. We characterized (i) CHCs and (ii) niches (through baiting, activity observations and foraging analysis). The species were distinct in both measures, although to a varying degree. Although they showed moderate niche partitioning along diet and environmental preferences, chemical differences were unexpectedly pronounced. This may be explained by divergent selection on mate recognition cues or by other influences on CHCs. Such striking chemical differences among closely related species may not be the rule and suggest that taxonomies based on CHCs should be interpreted cautiously; though, they remain useful tools for differentiating among cryptic species. [ABSTRACT FROM AUTHOR]

In animal societies, recognition of group members and relatives is an important trait for the evolution and maintenance of social behavior. In eusocial insects, nest mate recognition is based on cuticular hydrocarbons and allows colony members to reject competitors and parasites. The study of recognition cues in subsocial species can provide insights into evolutionary pathways leading to permanent sociality and kin-selected benefits of cooperation. In subsocial spiders, empirical evidence suggests the existence of both kin recognition and benefits of cooperating with kin, whereas the cues for kin recognition have yet to be identified. However, cuticular hydrocarbons have been proposed to be involved in regulation of tolerance and interattraction in spider sociality. Here, we show that subsocial Stegodyphus lineatus spiderlings have cuticular hydrocarbon profiles that are sibling-group specific, making cuticular hydrocarbons candidates for kin recognition cues. Our behavioral assays indicate that spiderlings can discriminate between cuticular cues from kin and nonkin: In a choice set-up, spiderlings more often chose to reside near cuticular chemical extracts of siblings compared with nonsiblings. Furthermore, we show that cuticular chemical composition changes during development, especially around the stage of dispersal, supporting the hypothesis that cuticular cues are involved in regulating conspecific tolerance levels. Lastly, our results indicate that the potential kin recognition cues might be branched alkanes that are influenced very little by rearing conditions and may be genetically determined. This indicates that a specific group of cuticular chemicals, namely branched alkanes, could have evolved to act as social recognition cues in both insects and spiders. [ABSTRACT FROM PUBLISHER]

Discrimination between nestmates and non-nestmates in social insects is thought to rely on the pattern of cuticular hydrocarbons. We investigated the cuticular hydrocarbon profiles of 2 parapatric sibling ant species, Temnothorax nylanderi (Förster, 1850) and Temnothorax crassispinus (Karavaiev, 1926), and their hybrid. We found that although the profiles show considerable similarities, a discriminant analysis based on the relative peak areas of cuticular hydrocarbons separates the 3 taxa. The profiles of hybrids were not consistently intermediate between those of the paternal species, suggesting either non-additive interactions among the parental biosynthetic pathways or systematic differences in environment-derived odor cues. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Social animals use recognition cues to discriminate between group members and non-members. These recognition cues may be conceptualized as a label, which is compared to a neural representation of acceptable cue combinations termed the template. In ants and other social insects, the label consists of a waxy layer of colony-specific hydrocarbons on the body surface. Genetic and environmental differences between colony members may confound recognition and social cohesion, so many species perform behaviors that homogenize the odor label, such as mouth-to-mouth feeding and allogrooming. Here, we test for another mechanism of cue exchange: indirect transfer of cuticular hydrocarbons via the nest material. Using a combination of chemical analysis and behavioral experiments with Camponotus aethiops ants, we show that nest soil indirectly transfers hydrocarbons between ants and affects recognition behavior. We also found evidence that olfactory cues on the nest soil influence nestmate recognition, but this effect was not observed in all colonies. These results demonstrate that cuticular hydrocarbons deposited on the nest soil are important in creating uniformity in the odor label and may also contribute to the template. [ABSTRACT FROM AUTHOR]

Background: In populations of most social insects, gene flow is maintained through mating between reproductive individuals from different colonies in periodic nuptial flights followed by dispersal of the fertilized foundresses. Some ant species, however, form large polygynous supercolonies, in which mating takes place within the maternal nest (intranidal mating) and fertilized queens disperse within or along the boundary of the supercolony, leading to supercolony growth (colony budding). As a consequence, gene flow is largely confined within supercolonies. Over time, such supercolonies may diverge genetically and, thus, also in recognition cues (cuticular hydrocarbons, CHC's) by a combination of genetic drift and accumulation of colony-specific, neutral mutations. Methodology/Principal Findings: We tested this hypothesis for six supercolonies of the invasive ant Anoplolepis gracilipes in north-east Borneo. Within supercolonies, workers from different nests tolerated each other, were closely related and showed highly similar CHC profiles. Between supercolonies, aggression ranged from tolerance to mortal encounters and was negatively correlated with relatedness and CHC profile similarity. Supercolonies were genetically and chemically distinct, with mutually aggressive supercolony pairs sharing only 33.1%±17.5% (mean ± SD) of their alleles across six microsatellite loci and 73.8%±11.6% of the compounds in their CHC profile. Moreover, the proportion of alleles that differed between supercolony pairs was positively correlated to the proportion of qualitatively different CHC compounds. These qualitatively differing CHC compounds were found across various substance classes including alkanes, alkenes and mono-, di- and trimethyl-branched alkanes. Conclusions: We conclude that positive feedback between genetic, chemical and behavioural traits may further enhance supercolony differentiation through genetic drift and neutral evolution, and may drive colonies towards different evolutionary pathways, possibly including speciation. [ABSTRACT FROM AUTHOR]

Background: Timely decision making is crucial for survival and reproduction. Organisms often face a speed-accuracy tradeoff, as fully informed, accurate decisions require time-consuming gathering and treatment of information. Optimal strategies for decision-making should therefore vary depending on the context. In mammals, there is mounting evidence that multiple systems of perceptual discrimination based on different neural circuits emphasize either fast responses or accurate treatment of stimuli depending on the context. Methodology/Principal Findings: We used the ant Camponotus aethiops to test the prediction that fast information processing achieved through direct neural pathways should be favored in situations where quick reactions are adaptive. Social insects discriminate readily between harmless group-members and dangerous strangers using easily accessible cuticular hydrocarbons as nestmate recognition cues. We show that i) tethered ants display rapid aggressive reactions upon presentation of non-nestmate odor (120 to 160 ms); ii) ants' aggressiveness towards non-nestmates can be specifically reduced by exposure to non-nestmate odor only, showing that social interactions are not required to alter responses towards non-nestmates; iii) decision-making by ants does not require information transfer between brain hemispheres, but relies on side-specific decision rules. Conclusions/Significance: Our results strongly suggest that first-order olfactory processing centers (up to the antennal lobes) are likely to play a key role in ant nestmate recognition. We hypothesize that the coarse level of discrimination achieved in the antennal lobes early in odor processing provides enough information to determine appropriate behavioral responses towards non-nestmates. This asks for a reappraisal of the mechanisms underlying social recognition in insects. [ABSTRACT FROM AUTHOR]

Social insects maintain colony cohesion by recognizing and, if necessary, discriminating against conspecifics that are not part of the colony. This recognition ability is encoded by a complex mixture of cuticular hydrocarbons (CHCs), although it is largely unclear how social insects interpret such a multifaceted signal. CHC profiles often contain several series of homologous hydrocarbons, possessing the same methyl branch position but differing in chain length (e.g., 15-methyl-pentatriacontane, 15-methyl-heptatriacontane, 15-methyl-nonatriacontane). Recent studies have revealed that within species these homologs can occur in correlated concentrations. In such cases, single compounds may convey the same information as the homologs. In this study, we used behavioral bioassays to explore how social insects perceive and interpret different hydrocarbons. We tested the aggressive response of Argentine ants, Linepithema humile, toward nest-mate CHC profiles that were augmented with one of eight synthetic hydrocarbons that differed in branch position, chain length, or both. We found that Argentine ants showed similar levels of aggression toward nest-mate CHC profiles augmented with compounds that had the same branch position but differed in chain length. Conversely, Argentine ants displayed different levels of aggression toward nest-mate CHC profiles augmented with compounds that had different branch positions but the same chain length. While this was true in almost all cases, one CHC we tested elicited a greater aggressive response than its homologs. Interestingly, this was the only compound that did not occur naturally in correlated concentrations with its homologs in CHC profiles. Combined, these data suggest that CHCs of a homologous series elicit the same aggressive response because they convey the same information, rather than Argentine ants being unable to discriminate between different homologs. This study contributes to our understanding of the chemical basis of nestmate recognition by showing that, similar to spoken language, the chemical language of social insects contains “synonyms,” chemicals that differ in structure, but not meaning. [ABSTRACT FROM AUTHOR]

Abstract  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. [ABSTRACT FROM AUTHOR]

Abstract  The cuticular hydrocarbons (CHCs) of the ant Lasius niger are described. We observe a high local colony specificity of the body cuticular profile as predicted for a monogynous and multicolonial species. The CHCs show a low geographical variation among different locations in France. The CHCs on the legs also are colony specific, but their relative quantities are slightly different from those on the main body. For the first time, we demonstrate that the inner walls of the ant nest are coated with the same hydrocarbons as those found on the cuticle but in different proportions. The high amount of inner-nest marking and its lack of colony-specificity may explain why alien ants are not rejected once they succeed in entering the nest. The cuticular hydrocarbons also are deposited in front of the nest entrance and on the foraging arena, with a progressive increase in n-alkanes relative amounts. Chemical marks laid over the substrate are colony specific only when we consider methyl-branched alkanes. Our data confirm that these “footprint hydrocarbons” are probably deposited passively by the contact of ant tarsae with the substrate. These results suggest that the CHCs chemical profiles used by ants in colony recognition are much more complex than a single template: ants have to learn and memorize odors that vary depending on their context of perception. [ABSTRACT FROM AUTHOR]

Background: Experiments under controlled laboratory conditions can produce decisive evidence for testing biological hypotheses, provided they are representative of the more complex natural conditions. However, whether this requirement is fulfilled is seldom tested explicitly. Here we provide a lab/field comparison to investigate the identity of an egg-marking signal of ant queens. Our study was based on ant workers resolving conflict over male production by destroying each other's eggs, but leaving queen eggs unharmed. For this, the workers need a proximate cue to discriminate between the two egg types. Earlier correlative evidence indicated that, in the ant Pachycondyla inversa, the hydrocarbon 3,11- dimethylheptacosane (3,11-diMeC27) is more abundant on the surface of queen-laid eggs. Methodology: We first tested the hypothesis that 3,11-diMeC27 functions as a queen egg-marking pheromone using laboratory-maintained colonies. We treated worker-laid eggs with synthetic 3,11-diMeC27 and found that they were significantly more accepted than sham-treated worker-laid eggs. However, we repeated the experiment with freshly collected field colonies and observed no effect of treating worker-laid eggs with 3,11-diMeC27, showing that this compound by itself is not the natural queen egg-marking pheromone. We subsequently investigated the overall differences of entire chemical profiles of eggs, and found that queen-laid eggs in field colonies are more distinct from worker-laid eggs than in lab colonies, have more variation in profiles, and have an excess of longer-chain hydrocarbons. Conclusions: Our results suggest that queen egg-marking signals are significantly affected by transfer to the laboratory, and that this change is possibly connected to reduced queen fertility as predicted by honest signaling theory. This change is reflected in the worker egg policing response under field and laboratory conditions. [ABSTRACT FROM AUTHOR]