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Olfactory systems in eusocial insects play a vital role in the discrimination of various chemical cues. Odorant receptors (ORs) are critical for odorant detection, and this family has undergone extensive expansion in ants. In this study, we re-annotated the OR genes from the most destructive invasive ant species Solenopsis invicta and 2 other Formicidae species, Ooceraea biroi and Monomorium pharaonis, with the aim of systematically comparing and analyzing the evolution and the functions of the ORs in ant species, identifying 356, 298, and 306 potential functional ORs, respectively. The evolutionary analysis of these ORs showed that ants had undergone chromosomal rearrangements and that tandem duplication may be the main contributor to the expansion of the OR gene family in S. invicta. Our further analysis revealed that 9-exon ORs had biased chromosome localization patterns in all three ant species and that a 9-exon OR cluster (SinvOR4–8) in S. invicta was under strong positive selection (Ka/Ks = 1.32). Moreover, we identified 5 S. invicta OR genes, namely SinvOR89, SinvOR102, SinvOR352, SinvOR327, and SinvOR135, with high sequence similarity (>70%) to the orthologs in O. biroi and M. pharaonis. An RT-PCR analysis was used to verify the antennal expression levels of these ORs, which showed caste-specific expression. The subsequent analysis of the antennal expression profiles of the ORs of the S. invicta workers from the polygyne and monogyne social forms indicated that SinvOR35 and SinvOR252 were expressed at much higher levels in the monogyne workers than in the polygyne workers and that SinvOR21 was expressed at higher levels in polygyne workers. Our study has contributed to the identification and analysis of the OR gene family in ants and expanded the understanding of the evolution and functions of the ORs in Formicidae species. [ABSTRACT FROM AUTHOR]

Abstract The ecological success of ants has made them abundant in most environments, yet inter‐ and intraspecific competition usually limit nest density for a given population. Most invasive ant populations circumvent this limitation through a supercolonial structure, eliminating intraspecific competition through a loss of nestmate recognition and lack of aggression toward non‐nestmates. Native to South America, Brachymyrmex patagonicus has recently invaded many locations worldwide, with invasive populations described as extremely large and dense. Yet, in contrast with most invasive ants, this species exhibits a multicolonial structure, whereby each colony occupies a single nest. Here, we investigated the interplay between genetic diversity, chemical recognition, and aggressive behaviors in an invasive population of B. patagonicus. We found that, in its invasive range, this species reaches a high nest density with individual colonies located every 2.5 m and that colony boundaries are maintained through aggression toward non‐nestmates. This recognition and antagonism toward non‐nestmates is mediated by chemical differentiation between colonies, as different colonies exhibit distinct chemical profiles. We highlighted that the level of aggression between colonies is correlated with their degree of genetic difference, but not their overall chemical differentiation. This may suggest that only a few chemical compounds influence nestmate recognition in this species or that weak chemical differences are sufficient to elicit aggression. Overall, this study demonstrates that invasive ant populations can reach high densities despite a multicolonial structure with strong aggression between colonies, raising questions about the factors underlying their ecological success and mitigating negative consequences of competitive interactions.

The benefits of cooperative living for foraging, nesting, defence and buffering environmental challenges lead animals with the most highly social lifestyles to dominate many ecosystems. However, living in larger, more highly connected groups should also increase the risks of pathogen exposure and transmission. While over long timescales selective responses could buffer the impacts of potential higher pathogen prevalence, similar processes are unlikely over short timescales. The red fire ant Solenopsis invicta is ideal for measuring the effects of group size on pathogen prevalence because two types of society coexist in this species: smaller single‐nest single‐queen colonies that are highly aggressive to their neighbours and larger multiple‐queen colonies that exchange resources with neighbouring nests. We compare the presence of viruses between these two colony types using metagenomic sequence classification of RNA‐sequencing reads. We find that queens from multiple‐queen colonies have 8.3‐times higher viral load and 1.5‐times higher viral diversity than queens from single‐queen colonies. This finding characterizes a rarely considered cost of transitions to more highly social living. Furthermore, our results show that highly social invertebrates can harbour many viruses. [ABSTRACT FROM AUTHOR]

Abstract Ants are among the most successful species at invading new environments. Their success undeniably comes from their various modes of reproduction and colony breeding structures, which influence their dispersal ability, reproductive potential, and foraging strategies. Almost all invasive ant species studied so far form supercolonies, a dense network of interconnected nests comprising numerous queens, without aggression toward non‐nestmates. This strategy results in invasive colonies that are able to grow extremely fast and large while avoiding intraspecific competition, allowing them to monopolize environmental resources and outcompete native species. Here, we developed and used 10 microsatellite markers to investigate the population structure and breeding system of the dark rover ant Brachymyrmex patagonicus Mayr in its introduced range. We determined whether this species exhibits a supercolonial structure by assessing whether different nests belonged to the same genetic colony. We inferred its dispersal ability by investigating isolation by distance and estimated the numbers of queens per colonies and mating per queen through parent‐offspring inferences. We found that most of the colonies of B. patagonicus were comprised of a single nest, headed by a single queen. Each nest was distinct from one another, without isolation by distance, which suggests strong dispersal ability through nuptial flights. These features are commonly observed in noninvasive and native ant species, but they are surprising for a successful invasive ant, as they strongly differ from other invasive ants. Overall, we discuss how this seemingly unfavorable strategy for an invasive ant might favor the invasive success of the dark rover ant in the United States.

The ecological success of ants has made them abundant in most environments, yet inter‐ and intraspecific competition usually limit nest density for a given population. Most invasive ant populations circumvent this limitation through a supercolonial structure, eliminating intraspecific competition through a loss of nestmate recognition and lack of aggression toward non‐nestmates. Native to South America, Brachymyrmex patagonicus has recently invaded many locations worldwide, with invasive populations described as extremely large and dense. Yet, in contrast with most invasive ants, this species exhibits a multicolonial structure, whereby each colony occupies a single nest. Here, we investigated the interplay between genetic diversity, chemical recognition, and aggressive behaviors in an invasive population of B. patagonicus. We found that, in its invasive range, this species reaches a high nest density with individual colonies located every 2.5 m and that colony boundaries are maintained through aggression toward non‐nestmates. This recognition and antagonism toward non‐nestmates is mediated by chemical differentiation between colonies, as different colonies exhibit distinct chemical profiles. We highlighted that the level of aggression between colonies is correlated with their degree of genetic difference, but not their overall chemical differentiation. This may suggest that only a few chemical compounds influence nestmate recognition in this species or that weak chemical differences are sufficient to elicit aggression. Overall, this study demonstrates that invasive ant populations can reach high densities despite a multicolonial structure with strong aggression between colonies, raising questions about the factors underlying their ecological success and mitigating negative consequences of competitive interactions. [ABSTRACT FROM AUTHOR]

Ants are among the most successful species at invading new environments. Their success undeniably comes from their various modes of reproduction and colony breeding structures, which influence their dispersal ability, reproductive potential, and foraging strategies. Almost all invasive ant species studied so far form supercolonies, a dense network of interconnected nests comprising numerous queens, without aggression toward non‐nestmates. This strategy results in invasive colonies that are able to grow extremely fast and large while avoiding intraspecific competition, allowing them to monopolize environmental resources and outcompete native species. Here, we developed and used 10 microsatellite markers to investigate the population structure and breeding system of the dark rover ant Brachymyrmex patagonicus Mayr in its introduced range. We determined whether this species exhibits a supercolonial structure by assessing whether different nests belonged to the same genetic colony. We inferred its dispersal ability by investigating isolation by distance and estimated the numbers of queens per colonies and mating per queen through parent‐offspring inferences. We found that most of the colonies of B. patagonicus were comprised of a single nest, headed by a single queen. Each nest was distinct from one another, without isolation by distance, which suggests strong dispersal ability through nuptial flights. These features are commonly observed in noninvasive and native ant species, but they are surprising for a successful invasive ant, as they strongly differ from other invasive ants. Overall, we discuss how this seemingly unfavorable strategy for an invasive ant might favor the invasive success of the dark rover ant in the United States. [ABSTRACT FROM AUTHOR]

Random amplified polymorphic DNA (RAPD) markers were used to investigate the genetic relationship among nests of the carpenter ant, Camponotus rufipes, located in the same area. Five random oligodecamers were used to amplify DNA from 108 ant workers collected from six nests. A total of 47 RAPD markers were identified, which revealed low levels of genetic differentiation among nests (Fst = 0.00218) and a low average Shannon index (0.3727) among workers within nests. These results together suggest that the C. rufipes nest may be formed by a single, once-mated queen and that nests produced by queens that are genetically related tend to keep their nests in close proximity to one other.

A remarkable social polymorphism is controlled by a single Mendelian factor in the fire ant Solenopsis invicta. A genomic element marked by the gene Gp-9 determines whether workers tolerate one or many fertile queens in their colony. Gp-9 was recently shown to be part of a supergene with two nonrecombining variants, SB and Sb. SB/ SB and SB/Sb queens differ in how they initiate new colonies, and in many physiological traits, for example odour and maturation rate. To understand how a single genetic element can affect all these traits, we used a microarray to compare gene expression patterns between SB/ SB and SB/Sb queens of three different age classes: 1-day-old unmated queens, 11-day-old unmated queens and mated, fully reproductive queens collected from mature field colonies. The number of genes that were differentially expressed between SB/ SB and SB/Sb queens of the same age class was smallest in 1-day-old queens, maximal in 11-day-old queens and intermediate in reproductive queens. Gene ontology analysis showed that SB/ SB queens upregulate reproductive genes faster than SB/Sb queens. For all age classes, genes inside the supergene were overrepresented among the differentially expressed genes. Consistent with the hypothesized greater number of transposons in the Sb supergene, 13 transposon genes were upregulated in SB/Sb queens. Viral genes were also upregulated in SB/Sb mature queens, consistent with the known greater parasite load in colonies headed by SB/Sb queens compared with colonies headed by SB/ SB queens. Eighteen differentially expressed genes between reproductive queens were involved in chemical signalling. Our results suggest that many genes in the supergene are involved in regulating social organization and queen phenotypes in fire ants. [ABSTRACT FROM AUTHOR]

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Solenopsis invicta Buren (red imported fire ant) are invasive pests that have the capability of major destructive impacts on lifestyle, ecology and economy. Control of this species is dependent, in part, upon ability to estimate the potential spread from newly discovered nests. The potential for spread and the spread characteristics differ between monogyne and polygyne social forms. Prior to this study, differentiation of the two social forms in laboratory test samples commonly used a method involving restriction endonuclease digestion of an amplified Gp-9 fragment. Success of this assay is limited by the quality of DNA, which in the field-collected insects may be affected by temporary storage in unfavourable conditions. Here, we describe an alternative and highly objective assay based upon a high resolution melt technique following preamplification of a significantly shorter Gp-9 fragment than that required for restriction endonuclease digestion. We demonstrate the application of this assay to a S. invicta incursion in Queensland, Australia, using field samples from which DNA may be partially degraded. The reductions in hands-on requirements and overall duration of the assay underpin its suitability for high-throughput testing. [ABSTRACT FROM AUTHOR]

Red imported fire ant, Solenopsis invicta Buren, workers were sampled from 26 colonies in Virginia during the 2007-2008 time period. Polymerase chain reaction (PCR) assays were used to determine colony social form (monogyny or polygyny) by genotyping ants at the Gp- 9 locus. Twenty of the colonies (76.9%) were found to be polygyne. Multiplex PCR was also used to detect the presence of several organisms currently being used as biological control agents for fire ants in the U.S., including the microsporidian parasite Kneallhazia solenopsae and Pseudacteon spp . parasitioid decapitating phorid flies in the sampled colonies. Kneallhazia solenopsae was detected in 11 of 26 colonies (42%). In addition, Pseudacteon spp. flies were detected in 2 (7.7%) colonies. The sampled colonies were examined by reverse transcription PCR (RT-PCR) for the presence of Solenopsis invicta viruses -1 and -2. Results indicated that 5 colonies were infected with SINV-1 (19%) and none were infected with SINV-2. This study is the first to characterize the red imported fire ant infestation in Virginia and documents the presence of biological control agents in this area. [ABSTRACT FROM AUTHOR]

Aim Understanding the factors underlying the successful establishment of invasive ant species is critical for developing quarantine strategies to prevent additional invasions as well as for determining how such species overcome the selective pressures in invaded areas. Although several studies have revealed differences in the social organization and population genetics of invasive ants in their native and introduced ranges, few studies have considered the potential interactions between alternate social forms within newly-invaded areas simply because many invasive ants are characterized as polygyne or unicolonial. Location Taoyuan and Chiayi County, Taiwan. Methods Both social forms (polygyne and monogyne) of the red imported fire ant, Solenopsis invicta, occur in two separately invaded areas in Taiwan (Taoyuan and Chiayi). We employed intensive sampling methods and diagnostic polymerase chain reaction assays to determine the distribution of these two social forms in both infested areas in Taiwan. Results The distribution of social forms differs dramatically between the two infested areas, consistent with different invasion histories regarding the original make-up of individuals comprising the initial founder group. The Taoyuan population likely was colonized initially by ants of both social forms, with the subsequent spread characterized by continuous outward movement of the two forms, particularly the monogyne form. In contrast, the initial founders of the Chiayi population likely were of the polygyne social form only, suggesting the monogyne social form in this population appeared only recently and likely arose directly from polygyne queens converting to the alternate social form. Main conclusions Our results provide detailed insights into the invasion history of S. invicta in Taiwan and suggest that the distinct reproductive biology of the two social forms may have shaped the current distribution of S. invicta in these infested areas and that the dynamics of two forms may affect the long-term persistence and potential for spread of this pest ant species. [ABSTRACT FROM AUTHOR]

Abstract: Thelohania solenopsae is a pathogen of the red imported fire ant, Solenopsis invicta, which debilitates queens and eventually causes the demise of colonies. Reductions of infected field populations signify its potential usefulness as a biological control agent. Thelohania solenopsae can be transmitted by introducing infected brood into a colony. The social forms of the fire ant, that is, monogyny (single queen per colony) or polygyny (multiple queens per colony), are associated with different behaviors, such as territoriality, that affect the degree of intercolony brood transfer. T. solenopsae was found exclusively in polygyne colonies in Florida. Non-synchronous infections of queens and transovarial transmission favor the persistence and probability of detecting infections in polygynous colonies. However, queens or alates with the monogyne genotype can be infected, and infections in monogyne field colonies have been reported from Louisiana and Argentina. Limited independent colony-founding capability and shorter dispersal of alate queens with the polygyne genotype relative to monogyne alates may facilitate the maintenance of infections in local polygynous populations. Demise of infected monogyne colonies can be twice as fast as in polygyne colonies and favors the pathogen’s persistence in polygyne fire ant populations. The social form of the fire ant reflects different physiological and behavioral aspects of the queen and colony that will impact T. solenopsae spread and ultimate usefulness for biological control. [Copyright &y& Elsevier]

Abstract: This is the first report of Thelohania solenopsae infections in monogyne (single-queen) Solenopsis invicta colonies in the field. In a 0.2-ha plot near Baton Rouge, Louisiana, inter-colony prevalence was 63% infection in June, 1999, when the population was 100% monogyne. In February, 2000, 21% of 33 monogyne and 90% of 10 polygyne colonies were infected. By May, 2001, the polygyne colonies had disappeared and only one of 34 monogyne colonies was infected, the final detection of T. solenopsae in the plot. Colony size did not differ significantly among the four types (monogyne versus polygyne×infected versus uninfected). [Copyright &y& Elsevier]

Changes in intraspecific social organization are of particular interest to sociobiologists, because an understanding of the genetic and ecological determinants of different social forms bears on the origins of sociality and mating isolation. The Southern Yellowjacket, Vespula squamosa (Drury), occurs as both polygyne and single queen nests. This study examined intra-caste size differences between both social forms of nests. Four polygyne and 13 single queen nests were collected from counties in Florida and Georgia. All single queen nests were one season old, whereas polygyne nests were at least 2 years old. The latter had significantly more workers per nest (p < 0.005), but mean values varied by over 50-fold. Workers and gynes from each social form were compared for six measures of body size: weight, wing length, femur length, thorax length, thorax width and head width. Correlation coefficients for pairs of all traits were highest between head width and both measures of thorax size (r > 0.72), whereas the lowest positive correlations were obtained between weight and all other measures of size (r < 0.64). Gynes of both social forms differed significantly in weight (p < 0.001) and thorax length (p < 0.04), whereas workers differed significantly for all six measures of size (p < 0.04 to p < 0.001). [ABSTRACT FROM AUTHOR]

This study reports previously undescribed behavior of fire ant queens ( Solenopsis invicta) on their nuptial flights. We captured large numbers of alate (winged) queens flying at low altitudes in dense swarms that were virtually devoid of males. We assayed the genotypes of these alate queens at the locus Gp-9, which exhibits strong genotype frequency differences between monogyne (single-queen) and polygyne (multiple-queen) populations, and found that almost all of these low-flying queens originated from polygyne colonies. Comparisons of mtDNA haplotype distributions of these queens to those of alates leaving polygyne nests suggest that the flying queens had not dispersed more than a few hundred meters. Moreover, the proportion of flying queens that were mated did not differ significantly from the proportion of reproductive queens that were mated within the same sites. Thus the flight behavior appears to occur subsequent to mating. We suggest that the flying queens are sampling the local environment in order to select a suitable landing site. Such a site would contain established polygyne nests into which the queens may be adopted as new reproductives. [ABSTRACT FROM AUTHOR]

1. Phorid flies in the genus Pseudacteon are parasitoids of ants. Variation in host size preferences of four South American and two North American Pseudacteon species on monogyne and polygyne forms of their host Solenopsis species [S. invicta Buren and S. geminata (F.), respectively] was documented. 2. Monogyne Solenopsis workers were, on average, significantly larger than polygyne workers, and the average size monogyne worker attacked was significantly larger than the average size polygyne worker attacked in four of the six Pseudacteon species. 3. Three South American Pseudacteon species attacked larger than average size workers, whereas one attacked smaller than average size workers, in both monogyne and polygyne forms. Both North American Pseudacteon species attacked larger than average size polygyne workers and smaller than average size monogyne workers. 4. Three Pseudacteon species were reared from eggs to adults in infected ants in the laboratory. The size of the emergent phorid fly was related positively to the size of the host worker ant, with females emerging from larger hosts. Similar patterns were documented for both monogyne and polygyne forms. 5. The mean size of worker host from which phorids emerged did not differ significantly between the monogyne and polygyne forms in the subsample of phorids reared to adults. 6. The observed patterns elucidate factors that may cause variation in Pseudacteon sex ratios, and have implications for biological control efforts of pest Solenopsis species. [ABSTRACT FROM AUTHOR]

Ants are among the most successful species at invading new environments. Their success undeniably comes from their various modes of reproduction and colony breeding structures, which influence their dispersal ability, reproductive potential, and foraging strategies. Almost all invasive ant species studied so far form supercolonies, a dense network of interconnected nests comprising numerous queens, without aggression toward non-nestmates. This strategy results in invasive colonies that are able to grow extremely fast and large while avoiding intraspecific competition, allowing them to monopolize environmental resources and outcompete native species. Here, we developed and used 10 microsatellite markers to investigate the population structure and breeding system of the dark rover ant Brachymyrmex patagonicus Mayr in its introduced range. We determined whether this species exhibits a supercolonial structure by assessing whether different nests belonged to the same genetic colony. We inferred its dispersal ability by investigating isolation by distance and estimated the numbers of queens per colonies and mating per queen through parent-offspring inferences. We found that most of the colonies of B. patagonicus were comprised of a single nest, headed by a single queen. Each nest was distinct from one another, without isolation by distance, which suggests strong dispersal ability through nuptial flights. These features are commonly observed in noninvasive and native ant species, but they are surprising for a successful invasive ant, as they strongly differ from other invasive ants. Overall, we discuss how this seemingly unfavorable strategy for an invasive ant might favor the invasive success of the dark rover ant in the United States., Competing Interests: None declared., (© 2019 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

The evolution of social life is usually associated with capabilities of individuals to protect group boundaries against foreign individuals. In colonies of ants, the number of reproductive queens is known to influence the accuracy of nestmate discrimination by resident workers. However, the pathway by which this effect is mediated remains unclear. The major hypothesis has long been that workers from multiple-queen colonies commit more discrimination errors against foreigners because their colonies contain a broader diversity of genetically determined cues characterising colony membership. Until recently, this hypothesis has received little attention and poor empirical support. In a recent study, Meunier et al.1 proposed an alternative, albeit not mutually exclusive hypothesis. The presence of one or multiple queens modifies chemical signals on colony members that trigger aggressive or cooperative behaviors during foreign encounters. Here, I detail how this new hypothesis is congruent with previous results and discuss potential limits and evolutionary implications of the two suggested hypotheses.