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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.)

Ants use a variety of semiochemicals for essential activities and have been a source for many novel natural products. While ant taxa produce a wide variety of chemicals, the chemistry and ecology of male ants have remained understudied. Tyramides are a class of compounds that have been found only in males of the Myrmicinae ant subfamily. Tyramides found in the fire ant Solenopsis invicta are transferred to gynes during mating where they are converted to tyramine, leading to rapid reproductive development. To further understand the evolution of tyramide production in male ants, we determined the tyramide composition in males of 15 fungus-growing ant species (Formicidae: Myrmicinae: Attini: Attina) and a Megalomyrmex species (Formicidae: Myrmicinae: Solenopsidini). Thirteen tyramides were identified, four for the first time in natural sources, and their percent composition was mapped to the fungus-growing ant phylogeny. [ABSTRACT FROM AUTHOR]

Chemical communication is common across all organisms. Insects in particular use predominantly chemical stimuli in assessing their environment and recognizing their social counterparts. One of the chemical stimuli used for recognition in social insects, such as ants, is the suite of long-chain, cuticular hydrocarbons. In addition to providing waterproofing, these surface hydrocarbons serve as a signature mixture, which ants can perceive, and use to distinguish between strangers and colony mates, and to determine caste, sex, and reproductive status of another individual. They can be both environmentally and endogenously acquired. The surface chemistry of adult workers has been studied extensively in ants, yet the pupal stage has rarely been considered. Here we characterized the surface chemistry of pupae of Formica exsecta, and examine differences among sexes, castes (reproductive vs. worker), and types of sample (developing individual vs. cocoon envelope). We found quantitative and qualitative differences among both castes and types of sample, but male and female reproductives did not differ in their surface chemistry. We also found that the pupal surface chemistry was more complex than that of adult workers in this species. These results improve our understanding of the information on which ants base recognition, and highlights the diversity of surface chemistry in social insects across developmental stages. [ABSTRACT FROM AUTHOR]

Queen pheromones evolved independently in multiple eusocial insect lineages, in which they mediate reproductive conflict by inhibiting worker ovarian development. Although fundamentally important for reproductive division of labor – the hallmark of eusociality – their evolutionary origins are enigmatic. Here, we analyze cuticular and Dufour's gland chemistries across alternative social and reproductive phenotypes in Megalopta genalis bees (tribe Augochlorini, family Halictidae) that facultatively express simple eusociality. Reproductive bees have distinct overall glandular and cuticular chemical phenotypes compared with non-reproductive workers. On the cuticle, a likely site of signal transmission, reproductives are enriched for certain alkenes, most linear alkanes, and are heavily enriched for all methyl-branched alkanes. Chemicals belonging to these compound classes are known to function as fertility signals in other eusocial insect taxa. Some macrocyclic lactones, compounds that serve as queen pheromones in the other eusocial halictid tribe (Halictini), are also enriched among reproductives relative to workers. The intra-population facultative eusociality of M. genalis permits direct comparisons between individuals expressing alternative reproductive phenotypes – females that reproduce alone (solitary reproductives) and social queens – to highlight traits in the latter that may be important mediators of eusociality. Compared with solitary reproductives, the cuticular chemistries of queens are more strongly differentiated from those of workers, and furthermore are especially enriched for methyl-branched alkanes. Determining the pheromonal function(s) and information content of the candidate signaling compounds we identify will help illuminate the early evolutionary history of queen pheromones, chemical signals central to the organization of insect eusocial behavior. [ABSTRACT FROM AUTHOR]

Ant queen pheromones (QPs) have long been known to affect colony functioning. In many species, QPs affect important reproductive functions such as diploid larvae sexualization and egg-laying by workers, unmated queens (gynes), or other queens. Until the 1990s, these effects were generally viewed to be the result of queen manipulation through the use of coercive or dishonest signals. However, in their seminal 1993 paper, Keller and Nonacs challenged this idea, suggesting that QPs had evolved as honest signals that informed workers and other colony members of the queen’s presence and reproductive state. This paper has greatly influenced the study of ant QPs and inspired numerous attempts to identify fertility-related compounds and test their physiological and behavioral effects. In the present article, we review the literature on ant QPs in various contexts and pay special attention to the role of cuticular hydrocarbons (CHCs). Although the controversy generated by Keller and Nonacs’ (Anim Behav 45:787-794, 1993) paper is currently less intensively debated, there is still no clear evidence which allows the rejection of the queen control hypothesis in favor of the queen signal hypothesis. We argue that important questions remain regarding the mode of action of QPs, and their targets which may help understanding their evolution. [ABSTRACT FROM AUTHOR]

Recognition is a fundamental process on which all subsequent behaviors are based at every organizational level, from the gene up to the super-organism. At the whole organism level, visual recognition is the best understood. However, chemical communication is far more widespread than visual communication, but despite its importance is much less understood. Ants provide an excellent model system for chemical ecology studies as it is well established that compounds known as cuticular hydrocarbons (CHCs) are used as recognition cues in ants. Therefore, stable species-specific odors should exist, irrespective of geographic locality. We tested this hypothesis by comparing the CHC profiles of workers of twelve species of Myrmica ants from four countries across Europe, from Iberia to the Balkans and from the Mediterranean to Fennoscandia. CHCs remained qualitatively stable within each species, right down to the isomer level. Despite the morphological similarity that occurs within the genus Myrmica, their CHCs were highly diverse but remarkably species-specific and stable across wide geographical areas. This indicates a genetic mechanism under strong selection that produces these species-specific chemical profiles, despite each species encountering different environmental conditions across its range. [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]

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]

While chemical communication regulates individual behavior in a wide variety of species, these communication systems are most elaborated in insect societies. In these complex systems, pheromones produced by the reproductive individuals (queens) are critical in establishing and maintaining dominant reproductive status over hundreds to thousands of workers. The proximate and ultimate mechanisms by which these intricate pheromone communication systems evolved are largely unknown, though there has been much debate over whether queen pheromones function as a control mechanism or as an honest signal facilitating cooperation. Here, we summarize results from recent studies in honey bees, bumble bees, wasps, ants and termites. We further discuss evolutionary mechanisms by which queen pheromone communication systems may have evolved. Overall, these studies suggest that queen-worker pheromone communication is a multi-component, labile dialog between the castes, rather than a simple, fixed signal-response system. We also discuss future approaches that can shed light on the proximate and ultimate mechanisms that underlie these complex systems by focusing on the development of increasingly sophisticated genomic tools and their potential applications to examine the molecular mechanisms that regulate pheromone production and perception. [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]

Using short (8-day) and long-term (28-day) experiments, we examined the stability of 20-hydroxyecdysone (20E) and the dominant phytosterols synthesized from a pulse of [2-14C]mevalonic acid ([14C]MVA) in hydroponically grown spinach ( Spinacia oleracea). In the short-term experiment, plant dry mass and shoot 20E pools steadily increased. Root uptake of [14C]MVA resulted in the stable incorporation of 14C radiolabel into whole plant 20E pools, with no significant changes over time. Levels of free and saponifiable phytosterols increased in the shoots while 14C-labeled shoot phytosterols remained constant. Unexpectedly, both 14C-labeled and unlabeled pools of root phytosterols decreased over time. In the long-term experiment, plant dry mass and shoot 20E levels increased over time, while total 14C-labeled 20E pools remained constant. Both root and shoot phytosterol pools increased over time while the 14C incorporation in these pools remained constant. Together these experiments indicate that 20E in spinach is metabolically stabile and thus shares this characteristic with plant terpenoids of known defensive function. While little is known about phytosterol turnover in plants, our results suggest that phytosterols can indeed exist in a very dynamic state but may also be stable over time. [ABSTRACT FROM AUTHOR]

Some plant defenses are known to be rapidly induced following attack by phytophagous insects. Plant-produced insect molting hormones, termed phytoecdysteroids, are believed to aid plant resistance; however, their dynamics are poorly understood. Using spinach ( Spinacia oleracea) as a model system, we examined the inducibility of phytoecdysteroids, primarily 20-hydroxyecdysone (20E), in an effort to characterize potential interactions with herbivorous insects. Rapid phytochemical induction was investigated using damage treatments and applications of defense-related plant-signal analogs, specifically methyl jasmonate (MJ) and methyl salicylate (MSA). Within two days, mechanically damaged roots exhibited two to three fold increases in phytoecdysteroid concentrations. Four days after root damage, small increases in shoot levels were also detectable. Unlike roots, foliar 20E concentrations were unaltered over a range of shoot treatments including insect herbivory ( Spodoptera exigua), mechanical damage, and MJ applications. Additions of MJ (12.5–50 μg/liter) to the root systems of hydroponically grown plants stimulated accumulations of root phytoecdysteroids in a dose-dependent manner, similar in magnitude to the response induced by root damage. Under identical conditions, MSA did not affect the accumulation of 20E when added to the hydroponic solutions of undamaged plants. Moreover, MSA inhibited the induction of 20E in wounded roots, but did not interfere with the action of applied MJ. In contrast to mechanical damage, roots did not induce 20E levels when challenged with two different fungal pathogens ( Pythium aphanidermatum and Phytophthora capsici).We propose that wound-induced accumulations of 20E are generated in the roots, signaled via endogenous jasmonates, and may confer enhanced resistance against subterranean herbivorous insects. [ABSTRACT FROM AUTHOR]