A new colony structure of the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>) in Southern Europe

The Argentine ant (Linepithema humile, Mayr) is a highly invasive species that has successfully spread from its native range in South America across many zones of the globe. In Southern Europe, two continental supercolonies have been identified, the Catalonian supercolony and the main European supercolony spreading over 6,000 km. In Corsica, a French Mediterranean island, the Argentine ant has been present for 60 years. Here we compare patterns of intraspecific aggression and cuticular hydrocarbon profiles of Argentine ants in Corsica to three mainland European colonies. Chemical analyses reveal the existence of cuticular signature variations among the six study sites relative to a gradient of aggression. We find two distinct colony groups not belonging to the Catalonian supercolony, suggesting that the new population originates either (1) from an independent introduction event from the native range resulting in a third European supercolony, or (2), given the chemical proximity and the moderate level of aggression between the two groups, from an existing European population followed by a drift producing a division within the main European supercolony.     

A Disjunct Argentine Ant Metacolony in Macaronesia and Southwestern Europe

The Argentine ant, Linepithema humile, originally from South America, is now a major pest in many parts of the world with Mediterranean-like climates. Earlier research indicated that southwestern European L. humile populations are segregated into two distinct supercolonies, the ‘Main’ supercolony extending through Portugal, Spain, southern France, and Italy, and the ‘Catalonian’ supercolony in eastern Spain. Both supercolonies are unicolonial, with workers showing no aggression towards members of the same supercolony, but severe aggression towards members of the other supercolony. Here we evaluated the behavioral relationships among non-native L. humile populations on the Macaronesian islands of Madeira and the Azores and non-native populations in southwestern Europe. We conducted aggression assays among L. humile workers from Madeira, the Azores, and the two southwestern European supercolonies. We found no aggressive interactions among any combination of workers from Madeira, the Azores, and the Main supercolony. However, workers from Madeira and the Azores always fought aggressively with workers from the Catalonian supercolony. Thus, the populations of L. humile in Madeira and the Azores appear to be unicolonial and act as if they belong to the Main supercolony of southwestern Europe. This set of geographically separated but mutually compatible supercolonies, which we term a ‘metacolony,’ appears to descend from one supercolony. Historical evidence suggests that the first L. humile population in the greater Mediterranean region was established in Madeira and that propagules from the dominant supercolony in Madeira gave rise to the dominant supercolonies in to other parts of the region.     

Chemical Discrimination and Aggressiveness via Cuticular Hydrocarbons in a Supercolony-Forming Ant,Formica yessensis

Background

Territorial boundaries between conspecific social insect colonies are maintained through nestmate recognition systems. However, in supercolony-forming ants, which have developed an extraordinary social organization style known as unicoloniality, a single supercolony extends across large geographic distance. The underlying mechanism is considered to involve less frequent occurrence of intraspecific aggressive behaviors, while maintaining interspecific competition. Thus, we examined whether the supercolony-forming species, Formica yessensis has a nestmate recognition system similar to that of the multicolonial species, Camponotus japonicus with respect to the cuticular hydrocarbon-sensitive sensillum (CHC sensillum), which responds only to non-nestmate CHCs. We further investigated whether the sensory system reflects on the apparent reduced aggression between non-nestmates typical to unicolonial species.

Methodology/Principal Findings

F. yessensis constructs supercolonies comprising numerous nests and constitutes the largest supercolonies in Japan. We compared the within-colony or between-colonies’ (1) similarity in CHC profiles, the nestmate recognition cues, (2) levels of the CHC sensillar response, (3) levels of aggression between workers, as correlated with geographic distances between nests, and (4) their genetic relatedness. Workers from nests within the supercolony revealed a greater similarity of CHC profiles compared to workers from colonies outside it. Total response of the active CHC sensilla stimulated with conspecific alien CHCs did not increase as much as in case of C. japonicus, suggesting that discrimination of conspecific workers at the peripheral system is limited. It was particularly limited among workers within a supercolony, but was fully expressed for allospecific workers.

Conclusions/Significance

We demonstrate that chemical discrimination between nestmates and non-nestmates in F. yessensis was not clear cut, probably because this species has only subtle intraspecific differences in the CHC pattern that typify within a supercolony. Such an incomplete chemical discrimination via the CHC sensilla is thus an important factor contributing to decreased occurrence of intraspecific aggressive behavior especially within a supercolony.     

Intercontinental union of Argentine ants: behavioral relationships among introduced populations in Europe, North America, and Asia

Many invasive ants, including the Argentine ant Linepithema humile, form expansive supercolonies, within which intraspecific aggression is absent. The behavioral relationships among introduced Argentine ant populations at within-country or within-continent scales have been studied previously, but the behavioral relationships among intercontinental populations have not been examined. The present study investigated the levels of aggression among intercontinental Argentine ant populations by transporting live ants from Europe and California to Japan and conducting aggression tests against Japanese populations. Workers from the dominant supercolonies of Europe and California did not show aggressive behavior toward workers from the dominant supercolony of Japan, whereas they fought vigorously against workers from minor supercolonies. The three massive supercolonies, together with Argentine ants from Macaronesia, may be the largest non-aggressive unit formed by a social insect species in which intraspecific aggression exists. Absence or low levels of aggression at transcontinental scale, which may have derived from low genetic variation, may help introduced Argentine ants maintain expansive supercolonies. The lack of aggression implies possible frequent exchanges of individuals among the intercontinental populations mediated by human activities.     

Contact between supercolonies elevates aggression in Argentine ants

Complex recognition systems underlie the social organization of many organisms. In social insects the acceptance of other individuals as nestmates can involve a variety of different cues, but the relative importance of these cues can change in relation to the fitness costs of accepting or rejecting other individuals. In this study we investigate the mechanisms that underlie recognition behaviour in Argentine ants (Linepithema humile). Introduced populations of Argentine ants are characterized by a social structure known as unicoloniality where intraspecific aggression is absent over large distances resulting in the formation of expansive supercolonies. Recent research has identified sites where multiple, mutually aggressive supercolonies co-occur allowing an examination of Argentine ant behaviour at territorial boundaries. We found that workers from different supercolonies always interact aggressively with one another, but that neighbours from different colonies (i.e., workers from nests located in the immediate vicinity of territory borders) consistently exhibited higher levels of aggression compared to those displayed by non-neighbours from different colonies (i.e., workers from nests located far enough away from a territory border so that interactions are unlikely). This difference in the level of aggression displayed between neighbours and between non-neighbours from different supercolonies cannot be explained by differences in relatedness or genetic similarity. Instead our findings suggest that direct contact between mutually antagonistic colonies is sufficient to elevate aggression. A laboratory experiment in which we manipulated the extent to which colonies with no prior history of contact could interact with one another, revealed that aggression increased after colonies were permitted to interact, but dropped after connections between colonies were severed. Moreover, the mere presence of an aggressive supercolony was sufficient to elicit elevated aggression. Overall these patterns are opposite to the “dear enemy” phenomenon and could be the result of the intense territorial aggression exhibited by established supercolonies of this species. Received 8 January 2007; revised 27 March 2007; accepted 28 March 2007.     

Differences in behavioural traits among native and introduced colonies of an invasive ant

Identifying the factors that promote the success of biological invasions is a key pursuit in ecology. To date, the link between animal personality and invasiveness has rarely been studied. Here, we examined in the laboratory how Argentine ant populations from the species’ native and introduced ranges differed in a suite of behaviours related to species interactions and the use of space. We found correlations among specific behavioural traits that defined an explorative-aggressive syndrome. The Main “European” supercolony (introduced range) more readily explored novel environments, displayed more aggression, detected food resources more quickly, and occupied more space than the Catalonian supercolony (introduced range) and two other Argentine supercolonies (native range). The two native supercolonies also differed in their personalities; one harbouring the less invasive personality, while the other is intermediate between the two introduced supercolonies. Therefore, instead of a binary pattern, Argentine ant supercolonies display a behavioural continuum that is independent on their geographic origin (native/introduced ranges). Our results also suggest that variability in personality traits is correlated to differences in the ecological success of Argentine ant colonies. Differences in group personalities may facilitate the persistence and invasion of animals under novel selective pressures by promoting adaptive behaviours. We stress that the concept of animal personality should be taken into account when elucidating the mechanisms of invasiveness.     

Multilevel genetic analyses of two European supercolonies of the Argentine ant, Linepithema humile

Some ants have an extraordinary unicolonial social organization, whereby individuals mix freely among physically separated nests. Recently, it was shown that the European population of Linepithema humile consisted of two enormous unicolonial supercolonies. Workers of the same supercolony are never aggressive to each other. In contrast, aggressiveness is invariably high between workers from different supercolonies. Here we investigated whether gene flow occurs between two supercolonies. We identified a contact zone in which we sampled 46 nests. For each nest, aggression tests were conducted against workers from reference nests from both supercolonies. Workers were always very aggressive towards workers of one of the supercolonies but not to workers of the other. Thus, all nests could be clearly assigned to one of the two supercolonies. For 22 of the 46 nests, we genotyped 15-16 workers at five microsatellite loci. A four-level hierarchical analysis of variance revealed very strong genetic differentiation between the two supercolonies (F(SUPERCOLONY-TOTAL) = 0.541) and low differentiation between sectors (i.e. group of nests connected together with trails) within supercolonies (F(SECTOR-SUPERCOLONY) = 0.064). The very high differentiation between the two supercolonies indicates a lack of ongoing gene flow, a conclusion further bolstered by the finding that the two supercolonies share no common alleles at two of the five microsatellite loci. A Bayesian clustering method also revealed the occurrence of two distinct clusters. These clusters exactly match the grouping obtained by aggression tests. None of the 332 genotyped individuals were admixed despite the fact that some nests of the two supercolonies were separated by less than 30 m. These results demonstrate that the two supercolonies have completely separate gene pools.     

DYNAMICS AND GENETIC STRUCTURE OF ARGENTINE ANT SUPERCOLONIES IN THEIR NATIVE RANGE

Some introduced ant populations have an extraordinary social organization, called unicoloniality, whereby individuals mix freely within large supercolonies. We investigated whether this mode of social organization also exists in native populations of the Argentine ant Linepithema humile . Behavioral analyses revealed the presence of 11 supercolonies (width 1 to 515 m) over a 3-km transect. As in the introduced range, there was always strong aggression between but never within supercolonies. The genetic data were in perfect agreement with the behavioral tests, all nests being assigned to identical supercolonies with the different methods. There was strong genetic differentiation between supercolonies but no genetic differentiation among nests within supercolonies. We never found more than a single mitochondrial haplotype per supercolony, further supporting the view that supercolonies are closed breeding units. Genetic and chemical distances between supercolonies were positively correlated, but there were no other significant associations between geographic, genetic, chemical, and behavioral distances. A comparison of supercolonies sampled in 1999 and 2005 revealed a very high turnover, with about one-third of the supercolonies being replaced yearly. This dynamic is likely to involve strong competition between supercolonies and thus act as a potent selective force maintaining unicoloniality over evolutionary time.     

The role of cuticular hydrocarbons as chemical cues for nestmate recognition in the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>)

Argentine ants (Linepithema humile) in their native South American range, like most other ant species, form spatially restricted colonies that display high levels of aggression toward other such colonies. In their introduced range, Argentine ants are unicolonial and form massive supercolonies composed of numerous nests among which territorial boundaries are absent. Here we examine the role of cuticular hydrocarbons (CHCs) in nestmate recognition of this highly damaging invasive ant using three supercolonies from its introduced range. We conducted behavioral assays to test the response of Argentine ants to workers treated with colonymate or non-colonymate CHCs. Additionally, we quantified the amount of hydrocarbons transferred to individual ants and performed gas chromatography-mass spectrometry (GC/MS) to qualitatively characterize our manipulation of CHC profiles. The GC/MS data revealed marked differences in the hydrocarbon profiles across supercolonies and indicated that our treatment effectively masked the original chemical profile of the treated ants with the profile belonging to the foreign individuals. We found that individual workers treated with foreign CHCs were aggressively rejected by their colonymates and this behavior appears to be concentration-dependent: larger quantities of foreign CHCs triggered higher levels of aggression. Moreover, this response was not simply due to an increase in the amount of CHCs applied to the cuticle since treatment with high concentrations of nestmate CHCs did not trigger aggression.The results of this study bolster the findings of previous studies on social insects that have implicated CHCs as nestmate recognition cues and provide insight into the mechanisms of nestmate recognition in the invasive Argentine ant. Received 6 February 2007; revised 31 May and 27 July 2007; accepted 16 August 2007.     

Abandoning aggression but maintaining self-nonself discrimination as a first stage in ant supercolony formation

An ant supercolony is a very large entity with very many queens. Although normal colonies of small extent and few queens remain distinct, a supercolony is integrated harmoniously over a large area [1, 2]. The lack of aggression is advantageous: Aggression is costly, involving direct and indirect losses and recognition errors [3, 4]. Indeed, supercolonial ants are among the ecologically most successful organisms [5-7]. But how supercolonies arise remains mysterious [1, 2, 8]. Suggestions include that reduced within-colony relatedness or reduced self-nonself discrimination would foster supercolony formation [1, 2, 5, 7, 9-12]. However, one risks confusing correlation and causality in deducing the evolution from distinct colonies to supercolonies when observing established supercolonies. It might help to follow up observations of another lack of aggression, that between single-queened colonies in some ant species. We show that the single-queened Lasius austriacus lacks aggression between colonies and occasionally integrates workers across colonies but maintains high within-colony relatedness and self-nonself discrimination. Provided that the ecological framework permits, reduced aggression might prove adaptive for any ant colony irrespective of within-colony relatedness. Abandoning aggression while maintaining discrimination might be a first stage in supercolony formation. This adds to the emphasis of ecology as central to the evolution of cooperation in general [13].     

Global-scale analyses of chemical ecology and population genetics in the invasive Argentine ant

Ants are some of the most abundant and ecologically successful terrestrial organisms, and invasive ants rank among the most damaging invasive species. The Argentine ant is a particularly well-studied invader, in part because of the extreme social structure of introduced populations, known as unicoloniality. Unicolonial ants form geographically vast supercolonies, within which territorial behaviour and intraspecific aggression are absent. Because the extreme social structure of introduced populations arises from the widespread acceptance of conspecifics, understanding how this colonymate recognition occurs is key to explaining their success as invaders. Here, we present analyses of Argentine ant recognition cues (cuticular hydrocarbons) and population genetic characteristics from 25 sites across four continents and the Hawaiian Islands. By examining both hydrocarbon profiles and microsatellite genotypes in the same individual ants, we show that native and introduced populations differ in several respects. Both individual workers and groups of nestmates in the introduced range possess less diverse chemical profiles than ants in the native range. As previous studies have reported, we also find that introduced populations possess much lower levels of genetic diversity than populations in the native range. Interestingly, the largest supercolonies on several continents are strikingly similar to each other, suggesting that they arose from a shared introduction pathway. This high similarity suggests that these geographically far-flung ants may still recognize and accept each other as colonymates, thus representing distant nodes of a single, widely distributed supercolony. These findings shed light on the behaviour and sociality of these unicolonial invaders, and pose new questions about the history and origins of introduced populations.     

Societies Drifting Apart? Behavioural,Genetic and Chemical Differentiation between Supercolonies in the Yellow Crazy Ant Anoplolepis gracilipes

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.     

Limited dispersal and an unexpected aggression pattern in a native supercolonial ant

Understanding how social groups function requires studies on how individuals move across the landscape and interact with each other. Ant supercolonies are extreme cooperative units that may consist of thousands of interconnected nests, and their individuals cooperate over large spatial scales. However, the inner structure of suggested supercolonial (or unicolonial) societies has rarely been extensively studied using both genetic and behavioral analyses. We describe a dense supercolony‐like aggregation of more than 1,300 nests of the ant Formica (Coptoformica) pressilabris. We performed aggression assays and found that, while aggression levels were generally low, there was some aggression within the assumed supercolony. The occurrence of aggression increased with distance from the focal nest, in accordance with the genetically viscous population structure we observe by using 10 DNA microsatellite markers. However, the aggressive interactions do not follow any clear pattern that would allow specifying colony borders within the area. The genetic data indicate limited gene flow within and away from the supercolony. Our results show that a Formica supercolony is not necessarily a single unit but can be a more fluid mosaic of aggressive and amicable interactions instead, highlighting the need to study internest interactions in detail when describing supercolonies.     

Genetic structure, nestmate recognition and behaviour of two cryptic species of the invasive big-headed ant Pheidole megacephala

Background

Biological invasions are recognized as a major cause of biodiversity decline and have considerable impact on the economy and human health. The African big-headed ant Pheidole megacephala is considered one of the world''s most harmful invasive species.

Methodology/Principal Findings

To better understand its ecological and demographic features, we combined behavioural (aggression tests), chemical (quantitative and qualitative analyses of cuticular lipids) and genetic (mitochondrial divergence and polymorphism of DNA microsatellite markers) data obtained for eight populations in Cameroon. Molecular data revealed two cryptic species of P. megacephala, one inhabiting urban areas and the other rainforests. Urban populations belong to the same phylogenetic group than those introduced in Australia and in other parts of the world. Behavioural analyses show that the eight populations sampled make up four mutually aggressive supercolonies. The maximum distance between nests from the same supercolony was 49 km and the closest distance between two nests belonging to two different supercolonies was 46 m. The genetic data and chemical analyses confirmed the behavioural tests as all of the nests were correctly assigned to their supercolony. Genetic diversity appears significantly greater in Africa than in introduced populations in Australia; by contrast, urban and Australian populations are characterized by a higher chemical diversity than rainforest ones.

Conclusions/Significance

Overall, our study shows that populations of P. megacephala in Cameroon adopt a unicolonial social structure, like invasive populations in Australia. However, the size of the supercolonies appears several orders of magnitude smaller in Africa. This implies competition between African supercolonies and explains why they persist over evolutionary time scales.     

Genetic structure of native ant supercolonies varies in space and time

Ant supercolonies are the largest cooperative units known in nature. They consist of networks of interconnected nests with hundreds of reproductive queens, where individuals move freely between nests, cooperate across nest boundaries and show little aggression towards non‐nestmates. The combination of high queen numbers and free mixing of workers, queens and brood between nests results in extremely low nestmate relatedness. In such low‐relatedness societies, cooperative worker behaviour appears maladaptive because it may aid random individuals instead of relatives. Here, we provide a comprehensive picture of genetic substructure in supercolonies of the native wood ant Formica aquilonia using traditional population genetic as well as network analysis methods. Specifically, we test for spatial and temporal variation in genetic structure of different classes of individuals within supercolonies and analyse the role of worker movement in determining supercolony genetic networks. We find that relatedness within supercolonies is low but positive when viewed on a population level, which may be due to limited dispersal of individuals and/or ecological factors such as nest site limitation and competition against conspecifics. Genetic structure of supercolonies varied with both sample class and sampling time point, which indicates that mobility of individuals varies according to both caste and season and suggests that generalizing has to be carried out with caution in studies of supercolonial species. Overall, our analysis provides novel evidence that native wood ant supercolonies exhibit fine‐scale genetic substructure, which may explain the maintenance of cooperation in these low‐relatedness societies.     

When supercolonies collide: territorial aggression in an invasive and unicolonial social insect

Some species of ants possess an unusual form of social organization in which aggression among nests is absent. This type of social organization, called unicoloniality, has been studied in only a handful of species and its evolutionary origins remain unclear. To date, no study has examined behavioural and genetic patterns at points of contact between the massive supercolonies that characterize unicoloniality. Since interactions at territory boundaries influence the costs of aggression and the likelihood of gene flow, such data may illuminate how supercolonies are formed and maintained. Here we provide field data on intraspecific territoriality for a widespread and invasive unicolonial social insect, the Argentine ant (Linepithema humile). We observed abrupt and well-defined behavioural boundaries at 16 contact zones between three different pairs of supercolonies. We visited nine of these zones weekly during a six-month period and observed consistent and intense intercolony aggression that resulted in variable, but often large, levels of worker mortality. Microsatellite variation along six transects across territory borders showed that F(ST) values were lower within supercolonies (0.08 +/- 0.01 (mean +/- SE)) than between supercolonies (0.29 +/- 0.01) and that this disparity was especially strong right at territory borders, despite direct and prolonged contact between the supercolonies. Matrix correspondence tests confirmed that levels of aggression and genetic differentiation were significantly correlated, but no relationship existed between geographic distance and either intraspecific aggression or genetic differentiation. Patterns of F(ST) variation indicated high levels of gene flow within supercolonies, but little to no gene flow between them. Overall, these findings are inconsistent with a model of relaxed ecological constraints leading to colony fusion and suggest that environmentally derived cues are not the prime determined of nestmate recognition in field populations of Argentine ants.     

Genetic and behavioural evidence for a city-wide supercolony of the invasive Argentine ant Linepithema humile (Mayr) (Hymenoptera: Formicidae) in southeastern Australia

The success of invasive ants is frequently attributed to genetic and behavioural shifts in colony structure during or after introduction. The Argentine ant ( Linepithema humile ), a global invader, differs in colony genetic structure and behaviour between native populations in South America and introduced populations in Europe, Japan, New Zealand and North America. However, little is known about its colony structure in Australia. We investigated the genetic structure and behaviour of L. humile across Melbourne, Victoria by quantifying variation at four microsatellite loci and assaying intraspecific aggression at neighbourhood (30–200 m), fine (1–3.3 km) and regional (5–82 km) spatial scales. Hierarchical analyses across these scales revealed that most genetic variation occurred among workers within nests (∼98%). However, although low genetic differentiation occurred among workers between nests at the fine and regional scales (∼2%), negligible differentiation was detected among workers from neighbouring nests. Spatial genetic autocorrelation analysis confirmed that neighbouring nests were genetically more similar to each other. Lack of aggression within and across these scales supported the view that L. humile is unicolonial and forms a large supercolony across Melbourne. Comparisons of genetic structure of L. humile among single nests sampled from Adelaide, Brisbane, Hobart and Perth with Melbourne showed no greater levels of genetic differentiation or dissimilar spatial structure, suggesting an Australia-wide supercolony.     

Queen execution increases relatedness among workers of the invasive Argentine ant,Linepithema humile

Polygyny in social insects can greatly reduce within‐nest genetic relatedness. In polygynous ant species, potential rival queens in colonies with multiple queens are often executed by other queens, workers, or both. The Argentine ant, Linepithema humile, native to South America, forms a “supercolony” that is composed of a large number of nests and is considered to contribute to the ant's invasion success. Currently, four mutually antagonistic supercolonies are contiguously distributed within a small area of Japan. Here, we analyzed the genetic structure and relatedness within and among the four supercolonies using microsatellite markers to clarify how L. humile maintains its supercoloniality. The results of AMOVA and BASP, the F_ST values, and the existence of several private alleles indicated that the L. humile population in the Kobe area had a characteristic genetic structure. Within a given supercolony, there was significant genetic differentiation (F_ST) among workers collected in May and those collected in September. The significant deviation from Hardy–Weinberg equilibrium increased, and the relatedness among workers significantly increased from May to September in all supercolonies. This result suggested that the supercolonies replaced old queens with new ones during the reproductive season, thus supporting the plausibility of queen execution. From the perspective of kin selection, workers collectively eliminate queens, thereby increasing their own inclusive fitness. Restricted gene flow among supercolonies, together with mating with sib and queen execution, could help to maintain the unique social structure of L. humile, the distribution of which is expanding worldwide.     

Parametric and non-parametric masking of randomness in sequence alignments can be improved and leads to better resolved trees

Background

Ants typically distinguish nestmates from non-nestmates based on the perception of colony-specific chemicals, particularly cuticular hydrocarbons present on the surface of the ants' exoskeleton. These recognition cues are believed to play an important role in the formation of vast so-called supercolonies that have been described for some invasive ant species, but general conclusions about the role of these cues are hampered by only few species being studied. Here we use data on cuticular hydrocarbons, aggression and microsatellite genetic markers to investigate the interdependence of chemical recognition cues, genetic distance and nestmate discrimination in the pharaoh ant (Monomorium pharaonis), a widespread pest species, and ask whether introduced populations of this species are genetically differentiated and exhibit intraspecific aggression.

Results

Microsatellite analyses of a total of 35 colonies from four continents revealed extremely high levels of genetic differentiation between almost all colonies (F _ST = 0.751 ± 0.006 SE) and very low within-colony diversity. This implies that at least 34 and likely hundreds more independent lineages of this ant have spread worldwide. Aggression tests involving workers from 14 different colonies showed only low levels of aggression, even between colonies that were geographically and/or genetically very distant. Chemical analyses of groups of worker ants showed that all colonies had the same cuticular compounds, which varied only quantitatively among colonies. There was a positive correlation between geographical and genetic distance, but no other significant relationships were detected between aggression, chemical profile, genetic distance and geographical distance.

Conclusions

The pharaoh ant has a global invasion history of numerous independent introductions resulting in genetically highly differentiated colonies typically displaying surprisingly low levels of intraspecific aggression, a behaviour that may have evolved in the native range or by lineage selection in the introduced range.     

Interspecific Aggression and Resource Monopolization of the Invasive Ant Anoplolepis gracilipes in Malaysian Borneo

Invasions by introduced ant species can be ecologically destructive and affect a wide range of taxa, particularly native ants. Invasive ant species often numerically dominate ant communities and outperform native ant species in effective resource acquisition. Here, we describe interactions between the invasive ant Anoplolepis gracilipes (Smith) and resident ant species in disturbed habitats in NE Borneo. We measured interference competition abilities of A. gracilipes by performing arena bioassays between two A. gracilipes colonies and seven local ant species, and measured its effective resource competition at baits within supercolonies and at supercolony boundaries. Furthermore, we compared ant species diversity and composition at baits among (A) core areas of A. gracilipes supercolonies, (B) supercolony boundaries and (C) outside supercolonies. Anoplolepis gracilipes was behaviorally dominant over most ant species except Oecophylla smaragdina. Within supercolonies, A. gracilipes discovered all food baits first, and monopolized the vast majority throughout the course of the experiment. At supercolony boundaries, A. gracilipes discovered baits later than resident ant species, but subsequently monopolized half of the baits. Furthermore, the activity and diversity of the ant community within A. gracilipes supercolonies was lower than at its boundaries and outside supercolonies, and the ant communities differed significantly between infested and noninfested areas. Our study supports the hypothesis that successful establishment of A. gracilipes in anthropogenically disturbed habitats may negatively affect resident ant communities through high levels of direct interspecific aggression and almost complete monopolization of resources within high‐density supercolonies.