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The Argentine ant is one of the five worst invasive ants. Recently it has been shown that one of the main compounds of its pygidial gland, iridomyrmecin, is used as a venom against competitors and enemies. Here, we explore the variability in the quantities of iridomyrmecin of individual workers, along a range of locations pertaining to both its native and invasive ranges, in order to know whether its venom could have contributed to the differential invasion success of European supercolonies. We specifically compared the amount of iridomyrmecin among supercolonies in the native range and among three invasive supercolonies: the Main supercolony (the most extended worldwide), the Corsican and the Catalonian supercolonies (both with a restricted distribution in Europe). Our main result is that the variability of the iridomyrmecin is very high. Looking at mean values, we found that the amount of iridomyrmecin of the Main supercolony was the lowest while the highest corresponded to the Corsican supercolony, with the Catalonian and the native range supercolonies having intermediate values. However, variability in the values within each supercolony was similar between supercolonies. This suggests that the success of a given invasive supercolony may not be explained by higher quantities of this defensive compound. Our results open the way for exploring the connection between defensive compounds and the invasion success of this global invader.

Two of the world’s most invasive ants, Linepithema humile and Lasius neglectus, are destined to overlap in range as they continue to spread throughout Europe. Although L. humile arrived first, and is therefore more numerically abundant, L. neglectus is the more behaviorally dominant of the two. We performed lab trials to determine whether L. humile could use numerical abundance to overcome the behavioral dominance of L. neglectus and whether the ants’ behavioral patterns shifted when the species co-occurred. We found that L. neglectus was more aggressive when less abundant, whereas the opposite was true of L. humile. When L. neglectus was outnumbered, it employed aggressive behaviors, such as biting or chemical attacks, more frequently than L. humile; it also utilized a behavioral sequence that included mandible opening and biting. Our results for these species support the hypothesis that species modulate their behavior towards competitors, which facilitates the understanding of how multiple invasive ant species can co-occur in a given area. Moreover, our study shows that the co-occurrence of invasive species could result from the use of two strategies: (1) the Bourgeois strategy, in which aggressiveness changes based on numerical dominance and (2) the dear-enemy strategy, in which aggressiveness is reduced when competitors co-occur. Since these strategies may lead to territory partitioning, we suggest that the behavioral flexibility displayed by both species when they overlap may allow local co-occurrence and increase their likelihood of co-occurrence during their range expansion in Europe, which could have a negative cumulative impact on invaded areas.

The Argentine ant (Linepithema humile, Mayr) is a highly invasive species. Recently, several RNA viruses have been identified in samples from invasive Argentine ant colonies. Using quantitative PCR, we investigated variation in the levels of these viruses in the main European supercolony over the course of a year. We discovered that virus prevalence and amounts of viral RNA were affected by season and caste: ants had more virus types during warm versus cold months, and queens had more virus types and higher virus prevalence than did workers or males. This seasonal variation was largely due to the appearance of positive-strand RNA viruses in the summer and their subsequent disappearance in the winter. The prevalences of positive-strand RNA viruses were positively correlated with worker foraging activity. We hypothesise that during warmer months, ants are more active and more numerous and, as a result, they have more conspecific and heterospecific interactions that promote virus transmission.

One of the main traits of invasive ants is the formation of supercolonies, large networks of polygynous nests lacking intraspecific competition, which allows them to reach high densities that facilitate their spread. However, different supercolonies exhibit different success in expanding along the world. Here, we explore whether the main chemical defensive compound of the Argentine ant could play a role in the differential invasiveness of supercolonies. We assessed differences in the amount of iridomyrmecin among supercolonies in the native range and in three invasive supercolonies: the Main supercolony (the most extended worldwide), and the Corsican and the Catalonian supercolonies (both with a restricted local distribution in Europe). We found that even if the amount of iridomyrmecin varied greatly between invaded regions in the three supercolonies in Europe and the native supercolonies in South America, the differences did not seem related to the success of invasion. The amount of iridomyrmecin of the Main supercolony was the lowest while the highest corresponded to the Corsican supercolony, with the Catalonian having intermediate values. This suggests that the success of a given invasive supercolony may not be explained by higher quantities of this defensive compound. Alternatively, reducing iridomyrmecin quantities in the invasive range could lead to more investment in other fitness traits that increase the invader's competitive ability. Our results open the way for exploring the contribution of defensive compounds in the competitive ability and spread of this global invader.

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.

Competition among queens in polygynous societies may result in queen executions or conflicts over personal reproduction. Understanding the factors that mediate the executions of ant queens should provide insight into how queen numbers are regulated in polygynous insect societies. The Argentine ant is a widespread invasive species that displays secondary polygyny, and workers execute 90% of their nestmate queens each spring. In this study, we investigated: (1) whether ambient temperature, queen number, and protein deprivation have an effect on queen executions and (2) whether workers select the queens slated for execution based on their cuticular hydrocarbon (CHC) profiles. We found that the percentage of queens executed was positively correlated with temperature and queen number but that protein deprivation did not play a role. As for queen fate, the levels of some CHCs were higher in surviving queens. One of these CHCs is associated with queen productivity (i.e egg-laying rate and ovarian index) suggesting that workers execute the least productive queens. Our findings suggest that chemical cues related to fertility signaling may mediate queen executions in Argentine ants Tis work was fnancially supported by the Spanish Ministry of Science and Innovation and the European Regional Development Fund (CGL2010-16451)

Social parasitism, i.e. the parasitic dependence of a social species on another free-living social species, is one of the most intriguing phenomena in social insects. It has evolved to various levels, the most extreme form being inquiline social parasites which have lost the worker caste, and produce only male and female sexual offspring that are reared by the host worker force. The inquiline syndrome has been reported in 4 species within the ant genus Plagiolepis, in Europe. Whether inquiline social parasitism evolved once or multiple times within the genus remains however unknown. To address this question, we generated data for 5 inquiline social parasites - 3 species previously described and 2 unidentified species - and their free-living hosts from Europe, and we inferred their phylogenetic relationships. We tested Emery's rule, which predicts that inquiline social parasites and their hosts are close relatives. Our results show that inquiline parasitism evolved independently at least 5 times in the genus. Furthermore, we found that all inquilines were associated with one of the descendants of their most related free-living species, suggesting sympatric speciation is the main process leading to the emergence of the parasitic species, consistent with the stricter version of Emery's rule.

The Argentine ant (Linepithema humile) is a highly invasive pest, yet very little is known about its viruses. We analysed individual RNA-sequencing data from 48 Argentine ant queens to identify and characterisze their viruses. We discovered eight complete RNA virus genomes – all from different virus families – and one putative partial entomopoxvirus genome. Seven of the nine virus sequences were found from ant samples spanning 7 years, suggesting that these viruses may cause long-term infections within the super-colony. Although all nine viruses successfully infect Argentine ants, they have very different characteristics, such as genome organization, prevalence, loads, activation frequencies and rates of evolution. The eight RNA viruses constituted in total 23 different virus combinations which, based on statistical analysis, were non-random, suggesting that virus compatibility is a factor in infections. We also searched for virus sequences from New Zealand and Californian Argentine ant RNA-sequencing data and discovered that many of the viruses are found on different continents, yet some viruses are prevalent only in certain colonies. The viral loads described here most probably present a normal asymptomatic level of infection; nevertheless, detailed knowledge of Argentine ant viruses may enable the design of viral biocontrol methods against this pest.

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., This work was funded by the Fyssen Foundation (postdoctoral contract to OB), the Spanish Ministry of Economy and Competitiveness, and the European Regional Development Fund (project CGL2012-36181 to XC and RB).

Polydomy associated with unicoloniality is a common trait of invasive species. In the invasive Argentine ant, colonies are seasonally polydomous. Most follow a seasonal fission-fussion pattern: they disperse in the spring and summer and aggregate in the fall and winter. However, a small proportion of colonies do not migrate; instead, they inhabit permanent nesting sites. These colonies are large and highly polydomous. The aim of this study was to (1) search for differences in the fecundity of queens between mother colonies (large and permanent) and satellite colonies (small and temporal), (2) determine if queens in mother and satellite colonies have different diets to clarify if colony size influences social organization and queen feeding, and (3) examine if colony location relative to the invasion front results in differences in the queen's diet. Our results indicate that queens from mother nests are more fertile than queens from satellite nests and that colony location does not affect queen oviposition rate. Ovarian dissections suggest that differences in ovarian morphology are not responsible for the higher queen oviposition rate in mother vs. satellite nests, since there were no differences in the number and length of ovarioles in queens from the two types of colonies. In contrast, the higher δ15N values of queens from mother nests imply that greater carnivorous source intake accounts for the higher oviposition rates This work was supported by the Ministry of Science and Innovation of the Government of Spain, and EU ERDF funds (CGL2010-16451)

What forces structure ecological assemblages? A key limitation to general insights about assemblage structure is the availability of data that are collected at a small spatial grain (local assemblages) and a large spatial extent (global coverage). Here, we present published and unpublished data from 51 ,388 ant abundance and occurrence records of more than 2,693 species and 7,953 morphospecies from local assemblages collected at 4,212 locations around the world. Ants were selected because they are diverse and abundant globally, comprise a large fraction of animal biomass in most terrestrial communities, and are key contributors to a range of ecosystem functions. Data were collected between 1949 and 2014, and include, for each geo-referenced sampling site, both the identity of the ants collected and details of sampling design, habitat type, and degree of disturbance. The aim of compiling this data set was to provide comprehensive species abundance data in order to test relationships between assemblage structure and environmental and biogeographic factors. Data were collected using a variety of standardized methods, such as pitfall and Winkler traps, and will be valuable for studies investigating large-scale forces structuring local assemblages. Understanding such relationships is particularly critical under current rates of global change. We encourage authors holding additional data on systematically collected ant assemblages, especially those in dry and cold, and remote areas, to contact us and contribute their data to this growing data set.

Pine forests on the northern plateau of the Iberian Peninsula have a long history of use, exploitation, and management. Current management practices consist of selective logging with relatively short periods of time between logging events. The main objective of this study is to detect changes in ant assemblages in the short time periods between selective logging activities. Ants were sampled at 44 sites considering three grouping categories of time periods after the last timber extraction: short (8. years). After selective logging the number of ant species increases as the forest recovers. A look at the differences between the assemblages when the analysis shifts from the species-specific level to functional groups showed differences between the short and long categories. This indicates that in certain circumstances the functional groups may be more informative of the functional restructuring of the ant assemblages in a disturbed habitat. Ant species from three functional groups display significant indicative values (Opportunist, Hot Climate Specialists and Subordinate Camponotini) in the medium- and long-time-after-logging categories: Messor capitatus (HCS) for medium-time category sites; and Aphaenogaster iberica (Op), Camponotus cruentatus (SC) and, Cataglyphis velox (HCS) for long-time category sites. No indicator species were found for the early stages of recovery. This information may also be of interest to managers because it reduces the number of data elements of the recovery status of these forests, and can be translated into monitoring protocols. The continued exploitation of these forests leads to an ant fauna that reflects this change. The results show that ant assemblages need at least 8. years to recover since only after that much time there is an emergence of Subordinate Camponotinae, a behaviorally dominant and low stress-tolerant functional group. This selective logging maintains the relative diversity and structure of ant assemblages This work was supported by the Ministry of Science and Innovation of the Government of Spain, and EU ERDF funds (CGL2010-16451)

Data concerning the influence of temperature on a species' physiological parameters can be a useful tool for predicting its potential distribution range, but in the case of the Argentine ant, data based on its physiological needs are too scarce and incomplete to make accurate predictions of this type. In the present study, we offer new and complete data concerning the Argentine ant queen's oviposition rate under a wide range of temperatures in the laboratory. We analyzed the oviposition rate of the Argentine ant (Linepithema humile) at 12 experimental temperatures: 10, 12, 14, 16, 18, 21, 24, 26, 28, 30, 32 and 34 degrees C under monogynous conditions (one queen per nest) and three different polygynous conditions (two, four and eight queens per nest). We found that temperature affected their oviposition rate and that the effect was similar regardless of the number of queens in the nest. Egg laying was at its maximum at 28 degrees C, with variation in the upper and lower temperature limits at which oviposition took place depending on the degree of polygyny. Oviposition rates were negatively correlated with the number of queens in the nest. We also observed a marked variation in the oviposition rate of queens subjected to the same experimental conditions.

Many studies have focused on the impacts of climate change on biological assemblages, yet little is known about howclimate interacts with other major anthropogenic influences on biodiversity, such as habitat disturbance. Using a unique global database of 1128 local ant assemblages, we examined whether climate mediates the effects of habitat disturbance on assemblage structure at a global scale. Species richness and evenness were associated positively with temperature, and negatively with disturbance. However, the interaction among temperature, precipitation and disturbance shaped species richness and evenness. The effectwas manifested through a failure of species richness to increase substantially with temperature in transformed habitats at low precipitation. At low precipitation levels, evenness increased with temperature in undisturbed sites, peaked at medium temperatures in disturbed sites and remained low in transformed sites. In warmer climates with lower rainfall, the effects of increasing disturbance on species richness and evenness were akin to decreases in temperature of up to 98C. Anthropogenic disturbance and ongoing climate change may interact in complicated ways to shape the structure of assemblages, with hot, arid environments likely to be at greatest risk. © 2015 The Author(s) Published by the Royal Society. All rights reserved.

The dynamic in Argentine ant colonies varies seasonally, influenced by biotic and abiotic factors. In winter the spatial range of the colony is contracted in large formations (winter nests) containing a large number of queens and workers. Winter nests are the clue to the species' dispersion power and the invasion of new habitats. For this reason a yearly elimination of queens and workers in winter at the edge (front) of the invasion could be a useful tool for weakening the species' dispersion and therefore limiting its establishment in new areas. Here, we determined the spatial dynamics of the Argentine ant nests during 1 year, and we assessed the invasion management by means of manual removal of winter nests for two consecutive winters, determining its effects during the following 3 years. We mapped nests found in 18 plots divided into two groups: extirpated (with removal of nests) and non-extirpated (control), along the fronts of three locations. Seasonal variation in the abundance of nests and workers, together with the two-year extirpation effects were evaluated. We found that colonies tended to follow an annual cycle of contraction and dispersion, with a decrease in the number of nests as we approached the invasion front. The extirpation was effective only in the front area, where it promoted smaller, less lasting and aggregated nests, as well as a decrease in the abundance of queens and workers. Nests also experienced a decrease during the two first winters but a recovery in the third, when no extirpation was done. Thus, a yearly perturbation should be performed to keep the expansion of the Argentine ant at a low rate, and to limit its establishment in new areas This work was financed by the Spanish Ministry of Science and Innovation with the support of a predoctoral grant (BES-2008-005102) associated with research projects MEC/FEDER2007-64080-C02-02/BOS and CGL2010-16451, and a FI grant from the European Social Fund and the DIUE of the Autonomous Government of Catalonia in support of MLE

Ants are good indicators of ecosystem health and therefore a good choice for rapid terrestrial bioassessments (RBAs) in land management. However, the application of these rapid protocols is unfeasible until efficient sampling methodologies adapted to management goals have been developed. Taking into account the need to improve tools to synthesize broad-scale RBA, the aim of this study is to search for a simple and efficient ant sampling protocol suitable to assess changes in ant assemblages taking, as an example, the specific case of the management of selective logging activities in public pine forests of the central Iberian Peninsula. Ants were sampled at eighteen sites. Each one corresponded to a . tranzón, a quadrangular unit of management. In our case all the . tranzones had similar areas (25-30. ha). Ants were sampled at each site using two methods: pitfall traps and hand collecting. Ant species richness (number of species) and the Shannon diversity index (H) were compared both for sampling method and for sample point. The Shannon diversity index was not considered as a biodiversity index, but a measure of entropy (uncertainty or information content) like in its original proposed sense. In both comparisons, hand collecting was the method that achieved a greater ant species collection and greater diversity indexes, and therefore, a higher information content, suggesting that it is the best choice to use for ant RBA protocols in the particular case of temperate forest habitats. This is an important issue for land managers of these forests in order to detect changes in ant assemblages between logging events. Moreover, the relatively large ant sizes reported in the Mohago pine forests in comparison with ant species across the whole Iberian Peninsula make the process of ant collection and recognition easier, simplifying forest management through the elaboration of simple protocols for rapid ant bioassessments (RBAs). Overall, this information greatly improves the current application of rapid ant assessment protocols for monitoring the recovery status of these temperate forests This work was supported by the Ministry of Science and Innovation of the Government of Spain, and EU ERDF funds (CGL2010-16451)

The Argentine ant is an invasive species that has been introduced worldwide causing devastating effects on entire ecosystems. Control strategies might be focused on slowing its rate of spread to limit its establishment inside yet non-invaded areas. For this, a better knowledge about nest selection is necessary to identify rapidly and accurately nest locations where to apply control measures. Herein, nest site selection by the Argentine ant, nests' physical characteristics and their longevity were studied in three invaded cork oak secondary forest. Results showed that this species shifts nest locations seasonally to keep appropriate microclimatic conditions, nesting mainly underneath rocks during cold and rainy months and in tree bases during warmer periods. The terrain features at micro-scale (orientation and slope) were found to influence the distribution of the Argentine ant nests beneath rocks. Additionally, artificial nests used as a control tool were tested, finding that their use may be suitable if they are set in appropriate locations and before the ants start migrating to winter aggregations Financial support was provided by the Spanish Ministry of Science and Education, with the associated projects CGL2007-64080-C02/BOS and CGL2010-16451/BOS, and the pre-doctoral grant BES-2008-372 005102 in support to MD

The influence of temperature on the developmental times and survival of insects can largely determine their distribution. For invasive species, like the Argentine ant, Linepithema humile Mayr (Hymenoptera: Formicidae), these data are essential for predicting their potential range based on mechanistic models. In the case of this species, such data are too scarce and incomplete to make accurate predictions based on its physiological needs. This research provides comprehensive new data about brood survival and developmental times at a wide range of temperatures under laboratory conditions. Temperature affected both the complete brood development from egg to adult worker and each of the immature stages separately. The higher the temperature, the shorter the development times. Brood survival from egg to adult was low, with the maximum survival rate being only 16% at 26 degrees C. Temperature also affected survival of each of the immature stages differently: eggs were negatively affected by high temperatures, while larvae were negatively affected by low temperatures, and the survival of pupae was apparently independent of environmental temperature. At 32 degrees C no eggs survived, while at 18 degrees C less than 2% of the eggs hatched into larva. The data from the present study are essential for developing prediction models about the distribution range of this tramp species based on its physiological needs in relation to temperature.

To address the lack of physiological approaches in current models assessing the potential distribution of the Argentine ant, we used data on brood development from distinct sources to evaluate a series of degree-day models for Catalonia (NE Iberian Peninsula), and data on the brood survival and oviposition rates to develop a worker production model. The degree-day model generated using data from Newell and Barber (1913) and Benois (1973) indicated that the number of degree-days required for the complete development from egg to adult worker was 445.4 degree-days above a threshold of 15.9 °C, while the model calibrated using data from Abril et al. (2008, in press) suggested 599.5 degree-days above 18.4 °C. Comparisons between the degree-day model predictions and the currently known distribution of the Argentine ant suggested that the one generated using data from Newell and Barber (1913) and Benois (1973) overestimated the presence of the species, while the one calibrated using data from Abril et al. (2008; in press) underestimated it. On the other hand, the predicted daily net production of Argentine ant workers generated by the worker production model predicted more accurately the distribution of the Argentine ant than the degree-day models. Our results show the utility of incorporating physiological data in models to assess the distribution limits of the Argentine ant, which up to date have taken little account of the physiological needs of the species in terms of its establishment and dispersion in its introduced ranges This study was financed by the Spanish Ministry of Education and Science (CGL2004-05240-C02-02/BOS and MEC/FEDER2007-64080-C02-02/BOS). S. Abril acknowledges support from the University of Girona, and N. Roura-Pascual from the Catalan Agency for Management of University and Research Grants (Generalitat de Catalunya) through a Beatriu de Pino´ s Postdoctoral Grant (2006 BPA 10124)

Background Reproductive division of labor in eusocial insects is a striking example of a shared genetic background giving rise to alternative phenotypes, namely queen and worker castes. Queen and worker phenotypes play major roles in the evolution of eusocial insects. Their behavior, morphology and physiology underpin many ecologically relevant colony-level traits, which evolved in parallel in multiple species. Results Using queen and worker transcriptomic data from 16 ant species we tested the hypothesis that conserved sets of genes are involved in ant reproductive division of labor. We further hypothesized that such sets of genes should also be involved in the parallel evolution of other key traits. We applied weighted gene co-expression network analysis, which clusters co-expressed genes into modules, whose expression levels can be summarized by their ‘eigengenes’. Eigengenes of most modules were correlated with phenotypic differentiation between queens and workers. Furthermore, eigengenes of some modules were correlated with repeated evolution of key phenotypes such as complete worker sterility, the number of queens per colony, and even invasiveness. Finally, connectivity and expression levels of genes within the co-expressed network were strongly associated with the strength of selection. Although caste-associated sets of genes evolve faster than non-caste-associated, we found no evidence for queen- or worker-associated co-expressed genes evolving faster than one another. Conclusions These results identify conserved functionally important genomic units that likely serve as building blocks of phenotypic innovation, and allow the remarkable breadth of parallel evolution seen in ants, and possibly other eusocial insects as well.