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.     

New mutualism for old: indirect disruption and direct facilitation of seed dispersal following Argentine ant invasion

The indirect effects of biological invasions on native communities are poorly understood. Disruption of native ant communities following invasion by the Argentine ant (Linepithema humile) is widely reported to lead indirectly to the near complete collapse of seed dispersal services. In coastal scrub in southeastern Australia, we examined seed dispersal and handling of two native and two invasive alien plant species at Argentine ant-invaded or -uninvaded sites. The Argentine ant virtually eliminates the native keystone disperser Rhytidoponera victoriae, but seed dispersal did not collapse following invasion. Indeed, Argentine ants directly accounted for 92% of all ant-seed interactions and sustained overall seed dispersal rates. Nevertheless, dispersal quantity and quality among seed species differed between Argentine ant-invaded and -uninvaded sites. Argentine ants removed significantly fewer native Acacia retinodes seeds, but significantly more small seeds of invasive Polygala myrtifolia than did native ants at uninvaded sites. They also handled significantly more large seeds of A. sophorae, but rarely moved them >5 cm, instead recruiting en masse, consuming elaiosomes piecemeal and burying seeds in situ. In contrast, Argentine ants transported and interred P. myrtifolia seeds in their shallow nests. Experiments with artificial diaspores that varied in diaspore and elaiosome masses, but kept seed morphology and elaiosome quality constant, showed that removal by L. humile depended on the interaction of seed size and percentage elaiosome reward. Small diaspores were frequently taken, independent of high or low elaiosome reward, but large artificial diaspores with high reward instead elicited mass recruitment by Argentine ants and were rarely moved. Thus, Argentine ants appear to favour some diaspore types and reject others based largely on diaspore size and percentage reward. Such variability in response indirectly reduces native seed dispersal and can directly facilitate the spread of an invasive alien shrub.     

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.     

Hydrocarbon-released nestmate aggression in the Argentine ant, Linepithema humile, following encounters with insect prey

Argentine ants, Linepithema humile, were attacked by their nestmates following contact with a particular prey item, the brown-banded cockroach, Supella longipalpa. Contact with prey, as brief as 2 min, provoked nestmate aggression. Argentine ants contaminated with hydrocarbons extracted from S. longipalpa also released nestmate aggression behavior similar to that released by the whole prey item, confirming the involvement of hydrocarbons. In contrast to S. longipalpa, little or no nestmate aggression was induced by other ant prey from diverse taxa. A comparison of prey hydrocarbon profiles revealed that all hydrocarbons of S. longipalpa were very long chain components with 33 or more carbons, while other prey had either less, or none, of the very long chain hydrocarbons of 33 carbons or greater. We identified the hydrocarbons of S. longipalpa and some new groups of long chain hydrocarbons of L. humile. The majority of S. longipalpa hydrocarbons were 35 and 37 carbons in length with one to three methyl branches, and closely resembled two previously unidentified groups of compounds from L. humile of similar chain length. The hydrocarbons of S. longipalpa and L. humile were compared and their role in the Argentine ant nestmate recognition is discussed.     

Consumption of introduced prey by native predators: Argentine ants and pit-building ant lions

When populations of native predators are subsidized by numerically dominant introduced species, the structure of food webs can be greatly altered. Surprisingly little is known, however, about the general factors that influence whether or not native predators consume introduced species. To learn more about this issue, we examined how native pit-building ant lions (Myrmeleon) are affected by Argentine ant (Linepithema humile) invasions in coastal southern California. Compared to areas without L. humile, invaded areas contained few native ant species and were deficient in medium-sized and large bodied native ants. Based on these differences, we predicted that Argentine ants would negatively affect ant lion larvae. Contrary to this expectation, observational surveys and laboratory growth rate experiments revealed that Myrmeleon were heavier, had longer mandibles, and grew more quickly when their main ant prey were Argentine ants rather than native ants. Moreover, a field transplant experiment indicated that growth rates and pupal weights were not statistically different for larval ant lions reared in invaded areas compared to those reared in uninvaded areas. Argentine ants were also highly susceptible to capture by larval Myrmeleon. The species-level traits that presumably make Argentine ant workers susceptible to capture by larval ant lions—small size and high activity levels—appear to be the same characteristics that make them unsuitable prey for vertebrate predators, such as horned lizards. These results underscore the difficulties in predicting whether or not numerically dominant introduced species serve as prey for native predators.     

Impacts of Argentine ants on avian nesting success

Biological invasions can have severe and widespread impacts on ecological communities. A few species of ants have become particularly damaging invaders but quantitative data of their impacts on many taxa is still lacking. We provide experimental evidence using artificial nests baited with quail eggs that the invasive Argentine ant (Linepithema humile) can be a significant avian nest predator – Argentine ants recruited to more nests and in higher abundance than the native ant species they displace. However, at a site invaded by Argentine ants, we monitored over 400 nests of a ground-nesting species, the Dark-eyed Junco (Junco hyemalis), and found that less than 2% of nests failed as a result of Argentine ant predation/infestation. A review of the literature also suggests that Argentine ants may not be a serious threat to bird nests relative to other predators or parasites. However, invasive ants with the capability of overwhelming prey though stinging (specifically the red-imported fire ant, Solenopsis invicta), may have a higher impact on avian nesting success. Received 14 January 2005; revised 28 April 2005; accepted 12 May 2005.     

Effects of habitat fragmentation on the three-way interaction among ants,aphids and larvae of the giant purple emperor, <Emphasis Type="Italic">Sasakia charonda</Emphasis> (Hewitson), a near-threatened butterfly

Survival rates of both early and middle instar larvae of the nymphalid butterfly, Sasakia charonda, were estimated to be lowest on test trees planted in a meadow (site A), intermediate in a small, narrow secondary deciduous broadleaf forest (small patch, site B) and highest in a large secondary deciduous broadleaf forest (large forest, site C). The larval mortality rates due to predation by tree-climbing predators from the ground (tree climbing predator) such as ants and the larvae of carabids were estimated to be greater at sites A and B than those at site C. The number of predatory ants climbing test trees was significantly greater at sites A and B than at site C, and the ants harvested honeydew from aphids living on tree leaves at those two sites. Aphid densities were significantly higher on trees at sites A and B than at site C, and aphid densities and numbers of predatory ants were significantly and positively correlated at sites A and B. In an experiment controlling aphid density per branch on test trees, the numbers of ants and the mortality rates of S. charonda larvae were greater on branches with high aphid densities than on those with low aphid densities at both sites A and B. These results suggest that the aphid density per host tree was higher in the meadow and the small patch than in the large forest; at both sites these higher aphid densities attracted higher numbers of predatory ants to test trees, and as a result, mortality rates of S. charonda larvae were increased.     

Nest raiding by the invasive Argentine ant on colonies of the harvester ant, Pogonomyrmex subnitidus

Invasive ants often displace native ants, and published studies that focus on these interactions usually emphasize interspecific competition for food resources as a key mechanism responsible for the demise of native ants. Although less well documented, nest raiding by invasive ants may also contribute to the extirpation of native ants. In coastal southern California, for example, invasive Argentine ants (Linepithema humile) commonly raid colonies of the harvester ant, Pogonomyrmex subnitidus. On a seasonal basis the frequency and intensity of raids vary, but raids occur only when abiotic conditions are suitable for both species. In the short term these organized attacks cause harvester ants to cease foraging and to plug their nest entrances. In unstaged, one-on-one interactions between P. subnitidus and L. humile workers, Argentine ants behaved aggressively in over two thirds of all pair-wise interactions, despite the much larger size of P. subnitidus. The short-term introduction of experimental Argentine ant colonies outside of P. subnitidus nest entrances stimulated behaviors similar to those observed in raids: P. subnitidus decreased its foraging activity and increased the number of nest entrance workers (many of which labored to plug their nest entrances). Raids are not likely to be the result of competition for food. As expected, P. subnitidus foraged primarily on plant material (85% of food items obtained from returning foragers), but also collected some dead insects (7% of food items). In buffet-style choice tests in which we offered Argentine ants food items obtained from P. subnitidus, L. humile only showed interest in dead insects. In other feeding trials L. humile consistently moved harvester ant brood into their nests (where they were presumably consumed) but showed little interest in freshly dead workers. The raiding behavior described here obscures the distinction between interspecific competition and predation, and may well play an important role in the displacement of native ants, especially those that are ecologically dissimilar to L. humile with respect to diet. Received 15 July 2005; revised 19 October 2005; accepted 26 October 2005.     

Interference competition by Argentine ants displaces native ants: implications for biotic resistance to invasion

The Argentine ant Linepithema humile (Dolichoderinae) is one of the most widespread invasive ant species in the world. Throughout its introduced range, it is associated with the loss or reduced abundance of native ant species. The mechanisms by which these native species are displaced have received limited attention, particularly in Australia. The role of interference competition in the displacement of native ant species by L. humile was examined in coastal vegetation in central Victoria (southeastern Australia). Foragers from laboratory colonies placed in the field consistently and rapidly displaced the tyrant ant Iridomyrmex bicknelli, the big-headed ant Pheidole sp. 2, and the pony ant Rhytidoponera victoriae from baits. Numerical and behavioural dominance enabled Argentine ants to displace these ants in just 20 min; the abundance of native species at baits declined 3.5–24 fold in direct relation to the rapid increase in L. humile. Most precipitous was the decline of I. bicknelli, even though species in this typically dominant genus have been hypothesized to limit invasion of L. humile in Australia. Interspecific aggression contributed strongly to the competitive success of Argentine ants at baits. Fighting occurred in 50–75% of all observed interactions between Argentine and native ants. This study indicates that Argentine ants recruit rapidly, numerically dominate, and aggressively displace from baits a range of Australian native ant species from different subfamilies and functional groups. Such direct displacement is likely to reduce native biodiversity and indirectly alter food web structure and ecosystem processes within invaded areas. Biotic resistance to Argentine ant invasion from native ants in this coastal community in southeastern Australia is not supported in this study.     

Rainfall facilitates the spread, and time alters the impact, of the invasive Argentine ant

Climate change may exacerbate invasions by making conditions more favorable to introduced species relative to native species. Here we used data obtained during a long-term biannual survey of the distribution of ant species in a 481-ha preserve in northern California to assess the influence of interannual variation in rainfall on the spread of invasive Argentine ants, Linepithema humile, and the displacement of native ant species. Since the survey began in 1993, Argentine ants have expanded their range into 74 new hectares. Many invaded hectares were later abandoned, so the range of Argentine ants increased in some years and decreased in others. Rainfall predicted both range expansion and interannual changes in the distribution of Argentine ants: high rainfall, particularly in summer months, promoted their spread in the summer. This suggests that an increase in rainfall will promote a wider distribution of Argentine ants and increase their spread into new areas in California. Surprisingly, the distribution of two native ant species also increased following high rainfall, but only in areas of the preserve that were invaded by L. humile. Rainfall did not have a negative impact on total native ant species richness in invaded areas. Instead, native ant species richness in invaded areas increased significantly over the 13 years of observation. This suggests that the impact of Argentine ants on naïve ant communities may be most severe early in the invasion process.     

Colony-structure variation and interspecific competitive ability in the invasive Argentine ant

The success of some invasive species may depend on phenotypic changes that occur following introduction. In Argentine ants ( Linepithema humile) introduced populations typically lack intraspecific aggression, but native populations display such behavior commonly. We employ three approaches to examine how this behavioral shift might influence interspecific competitive ability. In a laboratory experiment, we reared colonies of Forelius mccooki with pairs of Argentine ant colonies that either did or did not exhibit intraspecific aggression. F. mccooki reared with intraspecifically non-aggressive pairs of Argentine ants produced fewer eggs, foraged less actively, and supported fewer living workers than those reared with intraspecifically aggressive pairs. At natural contact zones between competing colonies of L. humile and F. mccooki, the introduction of experimental Argentine ant colonies that fought with conspecific field colonies caused L. humile to abandon baits in the presence of F. mccooki, whereas the introduction of colonies that did not fight with field colonies of Argentine ants resulted in L. humile retaining possession of baits. Additional evidence for the potential importance of colony- structure variation comes from the Argentine ants native range. At a site along the Rio de la Plata in Argentina, we found an inverse relationship between ant richness and density of L. humile (apparently a function of local differences in colony structure) in two different years of sampling.     

Impacts of the invasive Argentine ant on native ants and other invertebrates in coastal scrub in south‐eastern Australia

Invasive ants threaten native biodiversity and ecosystem function worldwide. Although their principal direct impact is usually the displacement of native ants, they may also affect other invertebrates. The Argentine ant, Linepithema humile (Dolichoderinae), one of the most widespread invasive ant species, has invaded native habitat where it abuts peri‐urban development in coastal Victoria in south‐eastern Australia. Here we infer impacts of the Argentine ant on native ants and other litter and ground‐dwelling invertebrates by comparing their abundance and taxonomic composition in coastal scrub forest either invaded or uninvaded by the Argentine ant. Species composition of native ants at bait stations and extracted from litter differed significantly between Argentine ant‐invaded and uninvaded sites and this was consistent across years. Argentine ants had a strong effect on epigeic ants, which were either displaced or reduced in abundance. The native ant Rhytidoponera victoriae (Ponerinae), numerically dominant at uninvaded sites, was completely absent from sites invaded by the Argentine ant. However, small hypogeic ants, including Solenopsis sp. (Myrmicinae) and Heteroponera imbellis (Heteroponerinae), were little affected. Linepithema humile had no detectable effect upon the abundance and richness of other litter invertebrates. However, invertebrate group composition differed significantly between invaded and uninvaded sites, owing to the varied response of several influential groups (e.g. Collembola and Acarina). Floristics, habitat structure and measured environmental factors did not differ significantly between sites either invaded or uninvaded by Argentine ants, supporting the contention that differences in native ant abundance and species composition are related to invasion. Changes in the native ant community wrought by Argentine ant invasion have important implications for invertebrate communities in southern Australia and may affect key processes, including seed dispersal.     

Protein marking reveals predation on termites by the woodland ant, <Emphasis Type="Italic">Aphaenogaster rudis</Emphasis>

Subterranean termites provide a major potential food source for forest-dwelling ants, yet the interactions between ants and termites are seldom investigated largely due to the cryptic nature of both the predator and the prey. We used protein marking (rabbit immunoglobin protein, IgG) and double antibody sandwich enzyme-linked immunosorbent assay (DAS-ELISA) to examine the trophic interactions between the woodland ant, Aphaenogaster rudis (Emery) and the eastern subterranean termite, Reticulitermes flavipes (Kollar). We marked the prey by feeding the termites paper treated with a solution of rabbit immunoglobin protein (IgG). Subsequently, we offered live, IgG-fed termites to ant colonies and monitored the intracolony distribution of IgG-marked prey. Laboratory experiments on the distribution of protein-marked termite prey in colonies of A. rudis revealed that all castes and developmental stages receive termite prey within 24 h. In field experiments, live, protein-marked termites were offered to foraging ants. Following predation, the marker was recovered from the ants, demonstrating that A. rudis preys on R. flavipes under field conditions. Our results provide a unique picture of the trophic-level interactions between predatory ants and subterranean termites. Furthermore, we show that protein markers are highly suitable to track trophic interactions between predators and prey, especially when observing elusive animals with cryptic food-web ecology. Received 19 January 2007; revised 23 March 2007; accepted 26 March 2007.     

Effects of Bt maize-fed prey on the generalist predator <Emphasis Type="Italic">Poecilus cupreus</Emphasis> L. (Coleoptera: Carabidae)

We investigated the effects of transgenic maize (Zea mays) expressing Bacillus thuringienses toxin (Bt maize) on larval and adult Poecilus cupreus carabid beetles in laboratory studies. In no-choice trials, neonate P. cupreus larvae were fed exclusively with Spodoptera littoralis caterpillars, which had been raised on Bt maize. S. littoralis raised on conventional maize or high quality Calliphora sp. pupae were fed to the beetle larvae in two control treatments. Bt-maize-fed caterpillar prey increased mortality to 100 within 40days. The experiment was repeated with 10-day-old beetle larvae. Bt treatment resulted in fewer pupae than in both controls, and in a higher mortality than in the Calliphora control. S. littoralis was suitable as exclusive prey in no-choice tests, at least for 40days, although prey quality seemed to be low compared to Calliphora pupae. The observed effects are most likely indirect effects due to further reduced nutritional prey quality. However, direct effects cannot be excluded. In the second part of the study, exposure of P. cupreus to Bt intoxicated prey was examined in paired-choice tests. Adult beetles were offered a choice between different prey conditions (frozen and thawed, freshly killed or living), prey types (S. littoralis caterpillars, Calliphorasp. pupae, cereal aphids) and prey treatments (raised on Bt or conventional maize). Living prey was preferred to frozen and dead prey. Caterpillars were only preferred to fly pupae and aphids when living. Prey treatment seemed to be least important for prey selection. The tests showed that P. cupreus ingested caterpillars readily and there was no evidence of them avoiding Bt containing prey, which means exposure in the field could occur. The presented protocols are a first step towards ecological risk assessment for carabid beetles.     

Foraging Ants as Scavengers on Entomopathogenic Nematode-Killed Insects

Ants were the most apparent invertebrate scavengers observed foraging on entomopathogenic nematode-killed insects (i.e., insect cadavers containing entomopathogenic nematodes and their symbiotic bacteria) in the present study. Workers of the Argentine ant,Linepithema humile(Mayr), scavenged nematode-killed insects on the surface and those buried 2 cm below the soil surface. Ant workers scavenged significantly more steinernematid-killed (60–85%) than heterorhabditid-killed (10–20%) insects. More 4-day-postinfected cadavers (hosts died within 48 h after exposure to nematodes) were scavenged than 10-day-postinfected cadavers. Ten-day-postinfected hosts contained live infective juvenile nematodes therefore ants may serve as phoretic agents. Other ant species, includingVeromessor andrei(Mayr),Pheidole vistanaForel,Formica pacificaFrancoeur, andMonomoriom ergatogynaWheeler, also scavenged nematode-killed insects. These ant species removed or destroyed about 45% of the steinernematid-killed insects. These results suggest that survival of steinernematid nematodes may be more significantly impacted by invertebrate scavengers, especially ants, than that of heterorhabditid nematodes, and placement of steinernematid-killed insects in the field for biological control may be an ineffective release strategy. Because entomopathogenic nematodes kill insects with the help of symbiotic bacteria, we tested the role of these bacterial species in deterring invertebrate scavengers by injecting bacteria (without nematodes) into insects and placing the cadavers in the field. None of the insects killed by the symbiotic bacterium,Photorhabdus luminescens(Thomas and Poinar) fromHeterorhabditis bacteriophoraPoinar, were scavanged, whereas 70% of the insects killed by the symbiotic bacterium,Xenorhabdus nematophilus(Poinar and Thomas) fromSteinernema carpocapsae(Weiser), and 90% of the insects killed byBacillus thuringiensisBerliner were scavenged by the Argentine ant. We conclude thatP. luminescensis responsible for preventing ants from foraging on heterorhabditid-killed hosts.     

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.     

Simple rules based on pile slope are used in the self organization of sand pile formation by <Emphasis Type="Italic">Pheidole oxyops</Emphasis> ants

We tested the hypothesis that slope influences where worker ants deposit excavated soil on piles near the nest entrance. We predicted that ants will deposit their load near the top of a pile where the slope changes from upward to downward, to prevent material rolling back towards the entrance. We tested this hypothesis by studying five natural colonies of Pheidole oxyops ants at a field site at S?o Sim?o, Brazil. At this site, each colony was dumping sandy soil excavated from its underground nest in a crescent-shaped pile c. 13 cm from and perpendicular to the nest entrance. Each nest was given an experimental sand pile of symmetrical curved cross section on a plywood platform that could be tilted 15 degrees up or down. From videos, the locations where individual ants dumped their soil loads were measured in relation to the inner (position = 0) and outer (position = 1) edges of the pile. When the platform was tilted down the ants deposited their loads significantly closer to the inner edge (0.458 ± 0.007) than when not tilted (0.530 ± 0.006). When the platform was tilted up the ants deposited their loads significantly further from the inner edge (0.626 ± 0.006) than when not tilted (0.522 ± 0.006). These results support the hypothesis that ants use pile slope in deciding where to dump their load. A similar rule is probably used in other ant species that place excavated soil in steep piles near the nest entrance. Received 5 February 2007; revised 10 June 2007; accepted 9 October 2007.     

Population genetic structure of the red imported fire ant, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>, in Taiwan

We generated and analyzed microsatellite genotypic data and mtDNA sequence data from the fire ant Solenopsis invicta collected from two separate infested areas (Taoyuan and Chiayi) in Taiwan to infer the population and colony structure of these recently established populations. These genetic analyses revealed the following patterns: 1) Relatedness among worker nestmates was significantly greater than zero for both social forms from both populations; 2) No significant isolation by distance was found among nests within each social form from either population; 3) Significant mtDNA but no nuclear differentiation occurs between sympatric social forms in Taoyuan; 4) Molecular signatures of genetic bottlenecks associated with recent introductions are evident in both populations; and 5) The two sampled populations, Taoyuan and Chiayi, are highly genetically differentiated at both the nuclear or mtDNA genomes and most likely derive from two separate introductions into Taiwan. While results from these analyses generally were consistent with predictions based on the known biology of these ants and similar studies of S. invicta in the U.S.A. and South America, some patterns likely reflect the recent introduction and human-mediated inadvertent transport of ants in Taiwan. This is the first study to investigate the population and colony structure of fire ants in Taiwan and results from our study represent an important contribution to the ongoing efforts aimed at eradicating this invasive pest in Taiwan. Received 18 June 2007; revised 11 October 2007; accepted 17 October 2007.     

Effect of Argentine ant invasions on ground-dwelling arthropods in northern California riparian woodlands

Although the Argentine ant (Linepithema humile) is a widespread invasive species that displaces native ants throughout its introduced range, the effects of these invasions on arthropods other than ants remain poorly known. This study documents the consequences of Argentine ant invasions on ants and other ground-dwelling arthropods in northern California riparian woodlands. Baits and unbaited pitfall traps were used to sample different components of the arthropod communities at five pairs of uninvaded and invaded sites. Sites occupied by Argentine ants supported almost no native epigeic ants except for the winter-active Prenolepis imparis. Sites with Argentine ants averaged four to ten times more ant workers than did sites with native ants, but ant worker biomass did not differ between uninvaded and invaded sites. Argentine ants recruited to baits in invaded areas, on average, in less than half the time of native ants in uninvaded areas. Despite the loss of epigeic native ants, higher Argentine ant worker abundance, and faster recruitment by Argentine ants at invaded sites, pitfall trap samples from uninvaded and invaded areas contained similar abundances and diversities of non-ant arthropods. These findings suggest that Argentine ants and the native ants they displace interact with the ground-dwelling arthropods of these habitats in a similar manner. Received: 24 February 1997 / Accepted: 9 November 1997     

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.