Kin structure and queen execution in the Argentine ant Linepithema humile

Every spring, workers of the Argentine Ant Linepithema humile kill a large proportion of queens within their nests. Although this behaviour inflicts a high energetic cost on the colonies, its biological significance has remained elusive so far. An earlier study showed that the probability of a queen being executed is not related to her weight, fecundity, or age. Here we test the hypothesis that workers collectively eliminate queens to which they are less related, thereby increasing their inclusive fitness. We found no evidence for this hypothesis. Workers of a nest were on average not significantly less related to executed queens than to surviving ones. Moreover, a population genetic analysis revealed that workers were not genetically differentiated between nests. This means that workers of a given nest are equally related to any queen in the population and that there can be no increase in average worker–queen relatedness by selective elimination of queens. Finally, our genetic analyses also showed that, in contrast to workers, queens were significantly genetically differentiated between nests and that there was significant isolation by distance for queens.     

Contrasting responses of native ant communities to invasion by an ant invader,Linepithema humile

Biological Invasions - Invasive alien species pose a serious threat to the integrity and function of natural ecosystems. Understanding how these invaders alter natural communities is therefore an...     

Genetic relationships among populations of the Argentine ant Linepithema humile introduced into Japan

Since some colonies of Argentine ant Linepithema humile were discovered in Japan in 1993, populations of this invasive alien ant species have been expanding their distribution. To resolve the number of invasions and the genetic structure in the early stages of introduction, we inferred the genetic structure and relationships among colonies from eight localities, from analyses of eight nuclear microsatellite DNA markers. F _ST analysis, principal component analysis and assignment test showed that at least three highly genetically differentiated groups of Argentine ants are present in Japan. Populations from Hiroshima Bay were grouped together (Hiroshima, Hatsukaichi, Otake, Iwakuni Central and Iwakuni Kuroiso), while those from Kobe and Yanai were both genetically distant from each other and from the Hiroshima Bay group. Hatsukaichi and Kobe are international seaports, suggesting that the by-ship invasion occurred at least twice. The invasion route of the Yanai population is unknown at this moment. The Aichi population was genetically distant from that of the Hiroshima Bay group by the difference in allele frequencies, and it was plausible that the Aichi population was introduced from the Hiroshima Bay group by human-mediated jump dispersal.     

Model of droplet dynamics in the Argentine ant Linepithema humile (Mayr)

The formation of droplets of ants Linepithema humile (Mayr) is observed under certain experimental conditions: a fluctuating aggregate forms at the end of a rod and a droplet containing up to 40 ants eventually falls down. When the flux of incoming ants is sufficient, this process can continue for several hours, leading to the formation and fall of tens of droplets. Previous work indicates that the time series of drop-to-drop intervals may result from a nonlinear low-dimensional dynamics, and the interdrop increments exhibit long-range anticorrelations. A model of aggregation and droplet formation, based on experimental observations, is introduced and shown to reproduce these properties.     

Loss of Wolbachia infection during colonisation in the invasive Argentine ant Linepithema humile

WOLBACHIA are maternally inherited bacteria, which are very common in arthropods and nematodes. Wolbachia infection may affect host reproduction through feminisation, parthenogenesis, male-killing, cytoplasmic incompatibility and increased fecundity. Previous studies showing discrepancies between the phylogenies of Wolbachia and its arthropod hosts indicate that infection is frequently lost, but the causes of symbiont extinction have so far remained elusive. Here, we report data showing that colonisation of new habitats is a possible mechanism leading to the loss of infection. The presence and prevalence of Wolbachia were studied in three native and eight introduced populations of the Argentine ant Linepithema humile. The screening shows that the symbiont is common in the three native L. humile populations analysed. In contrast, Wolbachia was detected in only one of the introduced populations. The loss of infection associated with colonisation of new habitats may result from drift (founder effect) or altered selection pressures in the new habitat. Furthermore, a molecular phylogeny based on sequences of the Wolbachia wsp gene indicates that L. humile has been infected by a single strain. Horizontal transmission of the symbiont may be important in ants as suggested by the sequence similarity of strains in the three genera Linepithema, Acromyrmex, and Solenopsis native from South and Central America.     

Effects of the Argentine ant Linepithema humile on seed dispersal and seedling emergence of Rhamnus alaternus

We studied influence of the Argentine ant's Linepithema humile occurrence on seed dispersal of Rhamnus alaternus (Rhamnaceae). Rhamnus alaternus is a fleshy fruit plant dispersed primarily by birds; the seeds have an elaiosome attractive to ants. The observations were made in two study plots of Mediterranean cork-oak forests (one invaded and the other not invaded by L. humile ) over two years. For R. alaternus , presence of L. humile was associated with of the following: reduction of seed transport to ant nests (80.2% of seed offered were removed in non-invaded vs 7.1% in invaded area), a shorter seed dispersal distance (mean=90.2 cm in non-invaded vs 1.1 cm in invaded area), increase in seed retention time on the soil surface (median time in non-invaded was 50 vs 209.2 min in invaded area) and increased vulnerability to predation. In addition there was lower probability of seedling emergence, due to little elaiosome removal from the seeds (82.0% of emergence for seeds without elaiosome and 57.3% for seeds with elaiosome). These results, similar to findings obtained in South African fynbos, confirm that the Argentine ant invasion can strongly affect ecosystem processes in the Mediterranean biome.     

Exploitation and interference competition between the invasive Argentine ant,Linepithema humile,and native ant species

Interactions between the invasive Argentine ant, Linepithema humile, and native ant species were studied in a 450-ha biological reserve in northern California. Along the edges of the invasion, the presence of Argentine ants significantly reduced the foraging success of native ant species, and vice versa. Argentine ants were consistently better than native ants at exploiting food sources: Argentine ants found and recruited to bait more consistently and in higher numbers than native ant species, and they foraged for longer periods throughout the day. Native ants and Argentine ants frequently fought when they recruited to the same bait, and native ant species were displaced from bait during 60% of these encounters. In introduction experiments, Argentine ants interfered with the foraging of native ant species, and prevented the establishment of new colonies of native ant species by preying upon winged native ant queens. The Argentine ants' range within the preserve expanded by 12 ha between May 1993 and May 1994, and 13 between September 1993 and September 1994, with a corresponding reduction of the range of native ant species. Although some native ants persist locally at the edges of the invasion of Argentine ants, most eventually disappear from invaded areas. Both interference and exploitation competition appear to be important in the displacement of native ant species from areas invaded by Argentine ants.     

The role of competitive dominance in the invasive ability of the Argentine ant (Linepithema humile)

To assess the importance of competition in the advance of invasive species, bait stations have been used to determine the dominance hierarchy of a community of native ants in Doñana National Park, southern Spain, and the status of the introduced species Linepithema humile (Argentine ant). Some native species, e.g. Cataglyphis floricola or Camponotus pilicornis, seem to be subordinate, i.e. occupy a low position in the competitive hierarchy; some are dominant (e.g. Pheidole pallidula), and others (e.g. Aphaenogaster senilis) occupy an intermediate position in the hierarchy. The Argentine ant is a competitively dominant species, because of its aggressive behavior and relative abundance. Irrespective of their position in the dominance hierarchy, L. humile and some native species adopt what games theory terms “the bourgeois strategy” during agonistic encounters with other species. Lone workers tend to be submissive in encounters whereas workers in the presence of other colony members are aggressive. L. humile was the only species which aggressively displaced large numbers of ants of other species from the bait. L. humile also expanded its range in the course of the experiment, displacing native species from parts of the study area.     

Chemical defense by the native winter ant (Prenolepis imparis) against the invasive Argentine ant (Linepithema humile)

The invasive Argentine ant (Linepithema humile) is established worldwide and displaces native ant species. In northern California, however, the native winter ant (Prenolepis imparis) persists in invaded areas. We found that in aggressive interactions between the two species, P. imparis employs a potent defensive secretion. Field observations were conducted at P. imparis nest sites both in the presence and absence of L. humile. These observations suggested and laboratory assays confirmed that P. imparis workers are more likely to secrete when outnumbered by L. humile. Workers of P. imparis were also more likely to secrete near their nest entrances than when foraging on trees. One-on-one laboratory trials showed that the P. imparis secretion is highly lethal to L. humile, causing 79% mortality. The nonpolar fraction of the secretion was chemically analyzed with gas chromatography/mass spectrometry, and found to be composed of long-chain and cyclic hydrocarbons. Chemical analysis of dissected P. imparis workers showed that the nonpolar fraction is derived from the Dufour's gland. Based on these conclusions, we hypothesize that this chemical defense may help P. imparis to resist displacement by L. humile.     

Acclimation effects on critical and lethal thermal limits of workers of the Argentine ant, Linepithema humile

For the Argentine ant Linepithema humile, bioclimatic models often predict narrower optimal temperature ranges than those suggested by behavioural and physiological studies. Although water balance characteristics of workers of this species have been thoroughly studied, gaps exist in current understanding of its thermal limits. We investigated critical thermal minima and maxima and upper and lower lethal limits following acclimation to four temperatures (15, 20, 25, 30 degrees C; 12L:12D photoperiod) in adult workers of the Argentine ant, L. humile, collected from Stellenbosch, South Africa. At an ecologically relevant rate of temperature change of 0.05 degrees Cmin(-1), CTMax varied between 38 and 40 degrees C, and CTMin varied between 0 and 0.8 degrees C. In both cases the response to acclimation was weak. A significant time by exposure temperature interaction was found for upper and lower lethal limits, with a more pronounced effect of acclimation at longer exposure durations. Upper lethal limits varied between 37 and 44 degrees C, whilst lower lethal limits varied between -4 and -10.5 degrees C, with an acclimation effect more pronounced for upper than lower lethal limits. A thermal envelope for workers of the Argentine ant is provided, demonstrating that upper thermal limits do likely contribute to distributional limits, but that lower lethal limits and limits to activity likely do not, or at least for workers who are not exposed simultaneously to the demands of load carriage and successful foraging behaviour.     

Osmolytes Contribute to pH Homeostasis of Escherichia coli

Background

Cytoplasmic pH homeostasis in Escherichia coli includes numerous mechanisms involving pH-dependent catabolism and ion fluxes. An important contributor is transmembrane K⁺ flux, but the actual basis of K⁺ compensation for pH stress remains unclear. Osmoprotection could mediate the pH protection afforded by K⁺ and other osmolytes.

Methods and Principal Findings

The cytoplasmic pH of E. coli K-12 strains was measured by GFPmut3 fluorimetry. The wild-type strain Frag1 was exposed to rapid external acidification by HCl addition. Recovery of cytoplasmic pH was enhanced equally by supplementation with NaCl, KCl, proline, or sucrose. A triple mutant strain TK2420 defective for the Kdp, Trk and Kup K⁺ uptake systems requires exogenous K⁺ for steady-state pH homeostasis and for recovery from sudden acid shift. The K⁺ requirement however was partly compensated by supplementation with NaCl, choline chloride, proline, or sucrose. Thus, the K⁺ requirement was mediated in part by osmolarity, possibly by relieving osmotic stress which interacts with pH stress. The rapid addition of KCl to strain TK2420 suspended at external pH 5.6 caused a transient decrease in cytoplasmic pH, followed by slow recovery to an elevated steady-state pH. In the presence of 150 mM KCl, however, rapid addition of another 150 mM KCl caused a transient increase in cytoplasmic pH. These transient effects may arise from secondary K⁺ fluxes occurring through other transport processes in the TK2420 strain.

Conclusions

Diverse osmolytes including NaCl, KCl, proline, or sucrose contribute to cytoplasmic pH homeostasis in E. coli, and increase the recovery from rapid acid shift. Osmolytes other than K⁺ restore partial pH homeostasis in a strain deleted for K⁺ transport.     

Absence of aggression but not nestmate recognition in an Australian population of the Argentine ant Linepithema humile

Intraspecific aggression is rare within introduced populations of the Argentine ant Linepithema humile, and colonies exhibit a structure known as unicoloniality, in which aggression among nests is atypical. We document a similar form of colony structure in an introduced population of Argentine ants in Victoria, Australia, in which aggression is extremely rare among nests ranging over hundreds of kilometres. However, using a highly sensitive behavioural bioassay we found that workers display subtle differences in their behaviour towards non-nestmates and nestmates. In particular, non-nestmates consistently engage in antennating behaviour with greater frequency than nestmates, perhaps providing a mechanism for homogenization of nest odour. Further, we found that non-nestmates at seaport sites (where populations may derive from multiple introductions) antennate each other with greater frequency than their counterparts from non-seaport sites. These data suggest that the Victorian population of L. humile may comprise multiple independent introductions. Received 4 July 2007; revised 15 January and 4 March 2008; accepted 4 March 2008.     

Trail Pheromone of the Argentine Ant,Linepithema humile (Mayr) (Hymenoptera: Formicidae)

The Argentine ant (Linepithema humile) is recognized as one of the world''s most damaging invasive species. One reason for the ecological dominance of introduced Argentine ant populations is their ability to dominate food and habitat resources through the rapid mobilization and recruitment of thousands of workers. More than 30 years ago, studies showed that (Z)-9-hexadecenal strongly attracted Argentine ant workers in a multi-choice olfactometer, suggesting that (Z)-9-hexadecenal might be the trail pheromone, or a component of a trail pheromone mixture. Since then, numerous studies have considered (Z)-9-hexadecenal as the key component of the Argentine ant trails. Here, we report the first chemical analyses of the trails laid by living Argentine ants and find that (Z)-9-hexadecenal is not present in a detectible quantity. Instead, two iridoids, dolichodial and iridomyrmecin, appear to be the primary chemical constituents of the trails. Laboratory choice tests confirmed that Argentine ants were attracted to artificial trails comprised of these two chemicals significantly more often than control trails. Although (Z)-9-hexadecenal was not detected in natural trails, supplementation of artificial dolichodial+iridomyrmecin trails with an extremely low concentraion of (Z)-9-hexadecenal did increase the efficacy of the trail-following behavior. In stark contrast with previous dogma, our study suggests that dolichodial and iridomyrmecin are major components of the Argentine ant trail pheromone. (Z)-9-hexadecenal may act in an additive manner with these iridoids, but it does not occur in detectable quantities in Argentine ant recruitment trails.     

Aggressive interactions between the introduced Argentine ant, Linepithema humile and the native odorous house ant, Tapinoma sessile

The Argentine ant (Linepithema humile) is an invasive species that disrupts the balance of natural ecosystems by displacing indigenous ant species throughout its introduced range. The mechanisms by which Argentine ants effectively compete against native ant species have been previously addressed in field studies that centered on interference and exploitation competition at baits and mainly examined the colony-level performance of Argentine ants. Detailed behavioral observations explaining the basis for the strong competitive ability of L. humile are comparatively rare. To gain a better understanding of the mechanisms by which Argentine ants displace native ants we examined the aggressive interactions between the Argentine ants and the odorous house ant, Tapinoma sessile in four different aggression assays: (1) worker dyad interactions, (2) symmetrical group interactions, (3) intruder introductions into an established resident colony, and (4) a resource competition assay which focused on competition for food and nesting space. Our results demonstrate a clear disparity between worker-level and colony-level fighting ability of Argentine ants and provide behavioral evidence to explain the superior interference ability of Argentine ants in group assays. Argentine ants experienced mixed success in fighting against odorous house ants in dyad interactions, but gradually gained a numerical advantage in symmetrical group interactions by active cooperation among nestmates. Results of the resource competition assay indicate that Argentine ants recruit rapidly, numerically dominate food and nesting sites, and aggressively displace T. sessile from baits. Taken together, the results of these assays allow us to pinpoint the behavioral mechanisms responsible for the remarkable competitive ability of Argentine ants.     

Long-term dynamics of the distribution of the invasive Argentine ant, Linepithema humile, and native ant taxa in northern California

Invasive species, where successful, can devastate native communities. We studied the dynamics of the invasive Argentine ant, Linepithema humile, for 7 years in Jasper Ridge, a biological preserve in northern California. We monitored the distributions at the hectare scale of native ant taxa and L. humile in the spring and fall from 1993 to 1999. We also studied the invasion dynamics at a finer resolution by searching for ants in 1-m² plots. Our results are similar at both scales. The distributions of several native species are not random with regard to L. humile; the distributions of several epigeic species with similar habitat affinities overlap much less frequently than expected with the distribution of L. humile. We found that season had a significant influence on the distributions of L. humile and several native taxa. Over the 7-year period, L. humile has increased its range size in Jasper Ridge largely at the expense of native taxa, but there is seasonal and yearly variation in this rate of increase. Studies of invasions in progress which sample across seasons and years may help to predict the spread and effects of invasive species.