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.     

Inferring historical introduction pathways with mitochondrial DNA: the case of introduced Argentine ants (Linepithema humile) into New Zealand

The threat imposed by invasive species and difficulties associated with control and management places more impetus on trying to prevent their introduction. The identification of introduction pathways is a vital component towards this goal. In this study, we use a genetic marker-based approach to retrospectively investigate the pathway of origin of the invasive Argentine ant ( Linepithema humile ) into New Zealand. We intensively sample the mitochondrial gene cytochrome b , from the entire known range of Argentine ants in New Zealand. No genetic variation was found in New Zealand. In order to identify likely introduction pathways, we use two alternative genetic analyses and suggest that a tcs approach that collapses identical haplotypes and calculates the probability of parsimony is superior to standard phylogenetic tree-building algorithms. A minimum spanning network allowed relationships to be examined among sequences collated from previous international studies. The cytochrome b sequence, when compared to a global database, matched that from an Australian population. That Australia is the potential source of Argentine ants is in agreement with the New Zealand interception record, as goods from Australia have the highest number of interception records of Argentine ants. Our approach can easily be duplicated for other organisms and the methodology can be more widely applied to help aid further efforts to identify the routes of transmission for other invasive species and allow us to efficiently direct our biosecurity monitoring effort.     

Relationships among native and introduced populations of the Argentine ant (Linepithema humile) and the source of introduced populations

The Argentine ant (Linepithema humile) is a damaging invasive species that has become established in many Mediterranean-type ecosystems worldwide. To identify likely sources of introduced populations we examined the relationships among native Linepithema populations from Argentina and Brazil and introduced populations of L. humile using mitochondrial cytochrome b sequence data and nuclear microsatellite allele frequencies. The mitochondrial phylogeny revealed that the populations in Brazil were only distantly related to both the introduced populations and the native populations in Argentina, and confirmed that populations in Brazil, previously identified as L. humile, are likely a different species. The microsatellite-based analysis provided resolution among native and introduced populations of L. humile that could not be resolved using the mitochondrial sequences. In the native range, colonies that were geographically close to one another tended to be genetically similar, whereas more distant colonies were genetically different. Most samples from the introduced range were genetically similar, although some exceptions were noted. Most introduced populations were similar to native populations from the southern Rio Parana and were particularly similar to a population from Rosario, Argentina. These findings implicate populations from the southern Rio Parana as the most likely source of introduced populations. Moreover, these data suggest that current efforts to identify natural enemies of the Argentine ant for biological control should focus on native populations in the southern Rio Parana watershed.     

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.     

Plasticity in queen number and social structure in the invasive Argentine ant (Linepithema humile)

In many polygynous social insect societies, ecological factors such as habitat saturation promote high queen numbers by increasing the cost of solitary breeding. If polygyny is associated with constrained environments, queen number in colonies of invasive social insects should increase as saturation of their new habitat increases. Here I describe the variation in queen number, nestmate relatedness, and nest size along a gradient of time since colonization in an invading population of Argentine ants (Linepithema humile) in Haleakala, Hawaii. Nest densities in this population increase with distance from the leading edge of the invasion, reaching a stable density plateau approximately 80 m from the edge (> 2 years after colonization). Although the number of queens per nest in Haleakala is generally lower than previously reported for Argentine ants, there is significant variation in queen number across this population. Both the observed and effective queen numbers increase across the density gradient, and nests in the center of the population contain queen numbers three to nine times higher than those on the edge of the invasion. The number of workers per nest is correlated with queen number, and nests in the center are six times larger than nests at the edge. Microsatellite analysis of relatedness among nestmates reveals that all nests in the Haleakala population are characterized by low relatedness and have evidence of multiple reproducing queens. Relatedness values are significantly lower in nests in the center of the population, indicating that the number of reproducing queens is greater in areas of high nest density. The variation in queen number and nestmate relatedness in this study is consistent with expectations based on changes in ecological constraints during the invasion of a new habitat, suggesting that the social structure of Argentine ant populations is strongly influenced by ecological factors. Flexibility in social structure may facilitate persistence in variable environments and may also confer significant advantages to a species when introduced into new areas.     

The Argentine ant, Linepithema humile, in Japan: Assessment of impact on species diversity of ant communities in urban environments

The Argentine ant Linepithema humile (Mayr) invaded the Hiroshima Prefecture in south‐west Japan some time before 1990. In this report, we describe the distribution of this exotic ant species and assess its impact on indigenous ant communities in urban areas. L. humile is now widely distributed mainly in urban areas and surrounding secondary vegetation of the cities Hatsukaichi and Hiroshima. The impact assessment suggested that L. humile reduced species diversity of local, indigenous ant communities. There was differential sensitivity of indigenous ant species to the invasion of L. humile. Some ant species disappeared in parks infested with L. humile; for example, Pheidole noda, Pheidole indica and Lasius japonicus. L. humile seemed to be superior to these ant species in certain traits and habits, such as mobility, recruitment ability, aggressiveness and omnivory. In contrast, Paratrechina sakurae and Camponotus vitiosus were less affected by L. humile infestation. The mechanisms allowing such coexistence seemed to be small body size (P. sakurae) and arboreal nesting habits (C. vitiosus).     

The Effect of Food Size and Dispersion Pattern on Retrieval Rate by the Argentine Ant, Linepithema humile (Hymenoptera: Formicidae)

Food acquisition in central-place foraging animals demands efficient detection and retrieval of resources. Most ant species rely on a mass recruitment foraging strategy, which requires that some potential foragers remain at the nest where they can be recruited to food once resources are found. Because this strategy reduces the number of workers initially looking for food, it may reduce the food detection rate while increasing the postdiscovery food retrieval rate. In previous studies this tradeoff has been analyzed by computer simulation and mathematical models. Both kinds of models show that food acquisition rate is greatly influenced by food distribution and resource patch size: as food is condensed into fewer patches, the maximal acquisition rate is achieved by a shift to fewer initial searchers and more potential recruits. In general, these models show that a mass recruitment strategy is most effective when resources are clumped. We tested this prediction in two experiments by letting laboratory colonies of the Argentine ant (Linepithema humile) forage for resources placed in different distributions. When all prey were small, retrieval rate increased with increasing resource patch size, in support of foraging models. When prey were large, however, the mass of prey returned to the colony over time was much lower than when prey were small and widely distributed. As more ants reached a large prey item, the distance the prey item was transported decreased due to a greater emphasis on feeding rather than transport. Because Argentine ants can transport more biomass externally than they can ingest, food retrieval that depends only on ingestion can depress the biomass retrieval rate. Thus, our results generally support theoretical foraging models, but we show how prey size, through differential prey-handling behavior, can produce an outcome greatly different from that predicted only on the distribution of resources.     

Effect of temperature on the oviposition rate of Argentine ant queens (Linepithema humile Mayr) under monogynous and polygynous experimental conditions

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.     

Parasitoids deter foraging by Argentine ants (Linepithema humile) in their native habitat in Brazil

The Argentine ant, Linepithema humile, has invaded sites across Africa, Australia, Europe, and North America. In its introduced ranges it eliminates native ants and tends agricultural pests. Few studies have examined the ecology of Argentine ants in their native habitat. This study examined the effects of parasitoid flies, genus Pseudacteon, on the foraging behavior of Argentine ants in part of their native range in southern Brazil. Pseudacteon parasitoids commonly attacked Argentine ants, but not other ant species, in daylight at temperatures above 18°C. Argentine ants abandoned food resources and returned underground in the presence of parasitoids. Parasitoid attack rates diminished as Argentine ants retreated underground. Where parasitoids were present, Argentine ants were abundant at food resources only during times of day when parasitoids were inactive. Where parasitoids were absent, Argentine ants were abundant at food resources throughout the day. Overall, the presence of parasitoids explained observed variation in Argentine ant foraging far better than temperature, although temperature had some effect. The results suggest that Pseudacteon parasitoids inhibit the ability of Argentine ants to gather food resources in their native habitat in Brazil. Received: 11 December 1997 / Accepted: 12 June 1998     

The Influence of Abiotic Factors and Temporal Variation on Local Invasion Patterns of the Argentine Ant (Linepithema humile)

The Argentine ant is a common pest of mediterranean-type ecosystems worldwide, causing widespread extirpation of many native ant species. This study examines spatial and temporal patterns of invasion at a local scale, investigates the effects of this invader on native ant communities and assesses causal mechanisms for these spatial, temporal and numerical patterns, particularly in relation to local climate. Argentine ant workers were more active in the drier months of summer and fall, and abundance was correlated with cumulative precipitation the previous winter. Argentine ant worker abundance greatly increased in conjunction with EI Niño events. No similar relationship existed for native ants. This invasive ant affected native ant abundances differently by species: some species were able to tolerate its presence temporarily. Overall, this study provides a framework of factors to consider when devising control strategies for the invasive Argentine ant.     

Let them eat termites—prey‐baiting provides effective control of Argentine ants,Linepithema humile,in a biodiversity hotspot

Invasive ants threaten biodiversity, ecosystem services and agricultural systems. This study evaluated a prey‐baiting approach for managing Argentine ants in natural habitat invaded by Argentine ants. Blackmound termites (Amitermes hastatus) were topically exposed to fipronil and presented to Argentine ants (Linepithema humile). In laboratory assays, L. humile colonies were offered fipronil‐treated termites within experimental arenas. The termites were readily consumed, and results demonstrate that a single termite topically treated with 590 ng fipronil is capable of killing at least 500 L. humile workers in 4 days. Field studies were conducted in natural areas invaded by L. humile. Fipronil‐treated termites scattered within experimental plots provided rapid control of L. humile and ant densities throughout the treated plots declined by 98 ± 5% within 21 days. Results demonstrate that the prey‐baiting approach is highly effective against L. humile and may offer an effective alternative to traditional bait treatments. Furthermore, prey‐baiting offers environmental benefits by delivering substantially less toxicant to the environment relative to current control methods which rely on commercial bait formulations and may offer greater target specificity.     

Analysis of whole-genome changes and characteristics of Linepithema humile (Hymenoptera: Formicidae) in the Republic of Korea

The Argentine ant, Linepithema humile (Mayr, 1868), native to the Paraguay River in South America, was first discovered in Korea in 2019. With increasing reports of L. humile, its genetic variation according to domestic growth colonies and its effects on the domestic ecosystem should be studied. Here, the genomes of L. humile specimens found at three locations in Busan were analyzed for genetic changes. First, morphological observation of L. humile samples collected from the three sites showed no phenotypic differences among them. Next, single-nucleotide polymorphism (SNP)/insertion and deletion (INDEL) analyses on the genomic DNA from the three groups showed that in the 1-NIE sample, the most frequent mutations were G → A and C → T. The mutations A → C and T → G were confirmed in the 14-NIE and 19-NIE samples. Although the number of SNPs in the N section was small, sequences of 4681 bp (1-NIE), 4217 bp (14-NIE) and 4631 bp (19-NIE) in length were identified. From the INDEL length distribution of the three samples, most changes were associated with insertions and deletions of 1–2 bp. However, no heterogeneity was found in the population samples analyzed based on SNP data. Comparative analysis of the SNPs investigated revealed that 760 819 (11.72%) of the total 6 492 517 SNPs were found in common, demonstrating that the three groups analyzed had different genetic backgrounds. Overall, we have developed a method for analyzing the genetic diversity of L. humile invading the Republic of Korea, precisely classified its genetic characteristics and obtained genomic data on interspecies mutations, according to the local environment.     

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.     

Geographical potential of Argentine ants (Linepithema humile Mayr) in the face of global climate change

Determining the spread and potential geographical distribution of invasive species is integral to making invasion biology a predictive science. We assembled a dataset of over 1000 occurrences of the Argentine ant (Linepithema humile), one of the world's worst invasive alien species. Native to central South America, Argentine ants are now found in many Mediterranean and subtropical climates around the world. We used this dataset to assess the species' potential geographical and ecological distribution, and to examine changes in its distributional potential associated with global climate change, using techniques for ecological niche modelling. Models developed were highly predictive of the species' overall range, including both the native distributional area and invaded areas worldwide. Despite its already widespread occurrence, L. humile has potential for further spread, with tropical coastal Africa and southeast Asia apparently vulnerable to invasion. Projecting ecological niche models onto four general circulation model scenarios of future (2050s) climates provided scenarios of the species' potential for distributional expansion with warming climates: generally, the species was predicted to retract its range in tropical regions, but to expand at higher latitude areas.     

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.     

Characterization of microsatellite loci for the Argentine ant,Linepithema humile,and their potential for analysis of colony structure in invading Hawaiian populations

We developed primers for five polymorphic microsatellite loci to analyse the genetic structure of colonies in an invading Argentine ant population located in Haleakala National Park on the island of Maui, Hawaii. Microsatellite loci were isolated using both a polymerase chain reaction‐based and a cloning‐based method. With a range of 3–18 alleles and expected levels of heterozygosity of 0.46–0.77, these loci provide useful markers for the detection of colony and population structure in new or expanding populations of this species.     

The effects of the invasive Argentine ant (Linepithema humile) and the native ant Prenolepis imparis on the structure of insect herbivore communities on willow trees (Salix lasiolepis)

Abstract 1. We examined the relative effects of the invasive Argentine ant, Linepithema humile, and a common native ant, Prenolepis imparis, on the community of herbivorous insects occurring on willow trees, Salix lasiolepis in Northern California, U.S.A. 2. Using paired control and treatment branches from which we excluded ants and other non‐volant predators, we found that effects of Argentine ants on the herbivore community were generally similar to those of P. imparis. Argentine ants and P. imparis suppressed the damage by skeletonising insects by 50%, but had little effect on most other external‐feeding or internal‐feeding guilds. 3. The abundance of aphids was 100% greater in the presence of Argentine ants, but there was no effect on aphid numbers in the presence of P. imparis. Late season aphid numbers were substantially higher in the presence of Argentine ants, but not P. imparis. 4. The effects of Argentine ants on skeletonising insects and aphids combined with the overwhelming abundance of Argentine ant workers, suggests that they may have substantial, but often overlooked, effects on the herbivore communities on other plant species in or near riparian habitats in which they invade.     

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.     

Exploitative strategies of the invasive Argentine ant (Linepithema humile) and native ant species in a southern Spanish pine forest

The Argentine ant, Linepithema humile (Mayr, 1868), is displacing native ant species in Do?ana National Park (Spain). This paper discusses the results of experiments aimed at analyzing exploitation competition between the invading species and other ant species in a park community. The Argentine ant was found to implement several strategies favoring its success in exploitation competition: mass recruitment, use of various microhabitats (on the ground and in trees), and activity over a wide range of temperatures. Although these strategies were not exclusive to L. humile, their joint use, together with the large number of workers forming each "unicolony," conferred a clear advantage for resource exploitation. Some native species were more severely affected than others by the presence of L. humile in terms of both abundance and behavior. The worst affected species were those whose ecological characteristics were similar to those of the Argentine ant, e.g., Pheidole pallidula (Nylander, 1849); the species least affected was Cataglyphis floricola Tinaut, 1993, possibly because of its subordinate and thermophilous nature (little overlap of daily activity rhythms with the exotic species).