Competitive Mechanisms Underlying the Displacement of Native Ants by the Invasive Argentine Ant

The Argentine ant (Linepithema humile) is a widespread invasive species that competitively displaces native ants throughout its introduced range. Although this pattern of displacement is well known, its underlying mechanisms remain little studied. To gain a more detailed understanding of this widespread competitive displacement, I compared the exploitative and interference abilities of the Argentine ant with those of seven species of native ants it displaces in riparian woodlands in northern California. I performed four different manipulative field experiments; each measured different aspects of the competitive ability of the eight species of ants in this study. The main goals of this study were to identify the mechanisms responsible for the Argentine ant’s strong competitive ability, to determine if native ants are subject to species-specific trade-offs in exploitative and interference ability typically present among coexisting ants, and if so, to assess whether Argentine ants are subject to this trade-off as well. Argentine ants located and recruited to baits as quickly or more quickly than did native ants—both in areas where Argentine ants and native ants occurred together (i.e., at the edge of invasion fronts) and where they occurred separately (i.e., away from invasion fronts). Along the edge of invasion fronts, Argentine ants also controlled a greater proportion of baits than did native ants. In one-on-one interactions, individual Argentine ant workers experienced mixed success in overcoming individual workers of the seven native ant species. When fighting against native ants, Argentine ants used both physical aggression and chemical defensive compounds, although the latter mechanism was more often successful in deterring opponents. Chemical defensive compounds produced by Argentine ants were repellent but appeared no more so than those of native ants. Although Argentine ant workers were not able to overcome native ant workers consistently, Argentine ant colonies succeeded in displacing most native ant colonies from baits. The discrepancy between worker-level and colony-level interference ability suggests that numerical advantages are key to the Argentine ant’s proficiency at interference competition. Like ants in other communities, the native ants in this study were subject to a competitive trade-off in which interference ability and exploitative ability were negatively correlated. In contrast, Argentine ants were proficient at both exploitative and interference competition relative to the native ants they displaced and are thus removed from this trade-off. These findings imply that Argentine ants secure a majority of available food resources where this species comes into contact with native ants. Argentine ants may be able to break the competitive trade-off constraining native ants because of their unique colony structure and because they have escaped their natural enemies. The observation that Argentine ants are uncoupled from the competitive trade-off constraining native ants may provide a general explanation for patterns of dominance within ant communities and for the success of other introduced species.

Ants are often considered antagonists when they visit flowers because they typically steal nectar without providing pollination services. Previous research on ant–flower interactions on two species of South African Proteaceae in the Cape Floral Kingdom revealed that the invasive Argentine ant (Linepithema humile), but not native ants, displace other floral arthropod visitors. To determine how common Argentine ant use of inflorescences is, how Argentine and native ant visits differ in the numbers they recruit to inflorescences, and what factors may affect Argentine and native ant foraging in inflorescences, I surveyed 723 inflorescences in 10 species in the genera Protea and Leucospermum across 16 sites and compared ant presence and abundance in inflorescences with abundance at nearby cat food and jam baits. Argentine ants were the most commonly encountered ant of the 22 observed. Argentine ants, as well as six species of native ants were present in all inflorescences for which they were present at nearby baits. Mean Argentine ant abundance per inflorescence was 4.4 ± 0.84 (SE) ants and similar to that of Anoplolepis custodiens and Crematogaster peringueyi, but higher than observed for the other most commonly encountered native ants, Camponotus niveosetosus and Lepisiota capensis. Both Argentine ants and A. custodiens were more likely to be found foraging in spring and under humid conditions, and in inflorescences closer to the ground, with lower sucrose concentrations, and with a greater proportion of open flowers. Argentine ants were more likely to be found in Protea inflorescences, whereas A. custodiens and L. capensis more often visited Leucospermum inflorescences. Considering its displacement of floral arthropods and widespread use of Proteaceae inflorescences, the Argentine ant could be posing a serious threat to plant and pollinator conservation in this biodiversity hotspot.

Invasive ants are notorious for directly displacing native ant species. Although such impacts are associated with Argentine ant invasions (Linepithema humile) worldwide, impacts within natural habitat are less widely reported, particularly those affecting arboreal ant communities. Argentine ants were detected in North Carolina mixed pine-hardwood forest for the first time but were localized on and around loblolly pines (Pinus taeda), probably because of association with honeydew-producing Hemiptera. We explored the potential impacts of L. humile on arboreal and ground-foraging native ant species by comparing interspersed loblolly pines invaded and uninvaded by Argentine ants. Impacts on native ants were assessed monthly over 1 yr by counting ants in foraging trails on pine trunks and in surrounding plots using a concentric arrangement of pitfall traps at 1, 2, and 3 m from the base of each tree. Of floristics and habitat variables, higher soil moisture in invaded plots was the only difference between plot types, increasing confidence that any ant community differences were caused by Argentine ants. Overall patterns of impact were weak. Composition differed significantly between Argentine ant invaded and uninvaded trunks and pitfalls but was driven only by the presence of Argentine ants rather than any resulting compositional change in native ant species. Native ant abundance and richness were similarly unaffected by L. humile. However, the abundance of individual ant species was more variable. Although numbers of the arboreal Crematogaster ashmeadi (Myrmicinae) declined on and around invaded pines, epigeic Aphaenogaster rudis (Myrmicinae) remained the most abundant species in all plots. Argentine ant densities peaked in late summer and fall, therefore overlapping with most native ants. Unexpected was their continued presence during even the coldest months. We provide evidence that Argentine ants can invade and persist in native North Carolina forests, probably mediated by pine-associated resources. However, their localized distribution and minimal impact on the native ant fauna relative to previously described invasions requires further resolution.

ABSTRACTArgentine ant (Linepithema humile (Mayr)) invasions are often associated with the displacement of ground‐dwelling arthropods. Argentine ant invasions can also exert other effects on the community through interactions with plants and their associated arthropods. For example, carbohydrate resources (e.g. floral or extrafloral nectar) may influence foraging behaviour and interactions among ants and other arthropods. In South Africa's Cape Floristic Region, Argentine ants and some native ant species are attracted to the floral nectar of Leucospermum conocarpodendron Rourke (Proteaceae), a native tree that also has extrafloral nectaries (EFNs). Despite having relatively low abundance in pitfall traps, Argentine ants visited inflorescences more frequently and in higher abundance than the most frequently observed native ants, Camponotus spp., though neither native nor Argentine ant floral foraging was influenced by the EFNs. Non‐metric multidimensional scaling revealed significant dissimilarity in arthropod communities on inflorescences with Argentine ants compared to inflorescences with native or no ants, with Coleoptera, Diptera, Hymenoptera, Arachnida, Orthoptera, and Blattaria all being underrepresented in inflorescences with Argentine ants compared to ant‐excluded inflorescences. Native honeybees (Apis mellifera capensis Eschscholtz) spent 75% less time foraging on inflorescences with Argentine ants than on inflorescences without ants. Neither Argentine ant nor native ant visits to inflorescences had a detectable effect on seed set of Le. conocarpodendron. However, a pollen supplementation experiment revealed that like many other proteas, Le. conocarpodendron is not pollen‐limited. Flower predation was negatively associated with increased ant visit frequency to the inflorescences, but did not differ among inflorescences visited by native and Argentine ants. Displacement of arthropods appears to be a consistent consequence of Argentine ant invasions. The displacement of floral arthropods by Argentine ants may have far‐reaching consequences for this biodiversity hotspot and other regions that are rich in insect‐pollinated plants.

The Argentine ant, Linepithema humile (Mayr), is a significant pest species, having become established on almost every continent, particularly in areas with a Mediterranean climate. In its introduced range, the Argentine ant has been associated with reductions in the abundance and diversity of native ant and nonant invertebrate fauna, as well as the interruption of ant–plant mutualisms. The distribution of Argentine ants has been correlated with particular abiotic factors, including soil moisture, relative humidity, disturbance, and particular soil and vegetation types. This study assessed the importance of temperature on the survival of Argentine ants and two native ant species (Iridomyrmex “rufoniger” and Rhytidoponera “convexa”) in the laboratory. Workers were placed in incubators of varying temperatures (25, 32, 40, 45, 47, and 50°C), and counts of the numbers dead and alive were recorded after 1, 2, and 3 h. The results showed that all species displayed almost 100% survival at 25, 32, 40, an...

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.

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.

We investigated the roles of habitat fragmentation and the invasion of an exotic species on the structure of ground-foraging ant communities in 40 scrub habitat fragments in coastal southern California. In particular, we asked: how do fragment age, fragment size, amount of urban edge, percentage of native vegetation, degree of isolation, and the relative abundance of an exotic species, the Argentine ant (Linepithema humile) affect native ants? Within these fragments, Argentine ants were more abundant near developed edges and in areas dominated by exotic vegetation. The number of native ground-foraging ant species at any point declined from an average of >7 to <2 species in the presence of the Argentine ant. Among fragments, a stepwise multiple regression revealed that the abundance of Argentine ants, the size of the fragment, and the number of years since it was isolated from larger continuous areas of scrub habitat best predict the number of remaining native ant species. The Argentine ant was found in every fragment surveyed as well as around the edges of larger unfragmented areas. Fragments had fewer native ant species than similar-sized plots within large unfragmented areas, and fragments with Argentine ant-free refugia had more native ant species than those without refugia. The relative vulnerability of native ants to habitat fragmentation and the subsequent presence of Argentine ants vary among species. The most sensitive species include army ants (Neivamyrmex spp.) and harvester ants (genera Messor and Pogonomyrmex), both of which are important to ecosystem-level processes. Our surveys suggest that the Argentine ant is widespread in fragmented coastal scrub habitats in southern California and strongly affects native ant communities.

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. Previous studies that examined the mechanisms by which Argentine ants attain ecological dominance showed that superior interference and exploitation competition are key to the successful displacement of native ant species. The objective of this research was to test the hypothesis that effective interference competition by Argentine ants may also be detrimental to the survival of Argentine ant colonies where Argentine ants and native ants compete at toxic baits used to slow the spread of Argentine ants. To study this hypothesis, we examined the competitive interactions between Argentine ants and native odorous house ants, Tapinoma sessile, in the presence and absence of toxic baits. Results showed that Argentine ants aggressively outcompete T. sessile from toxic baits through efficient interference competition and monopolize bait resources. This has severe negative consequences for the survival of Argentine ants as colonies succumb to the toxic effects of the bait. In turn, T. sessile avoid areas occupied by Argentine ants, give up baits, and consequently suffer minimal mortality. Our results provide experimental evidence that highly efficient interference competition may have negative consequences for Argentine ants in areas where toxic baits are used and may provide a basis for designing innovative management programs for Argentine ants. Such programs would have the double benefit of selectively eliminating the invasive species while simultaneously protecting native ants from the toxic effects of baits.

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.

Biological invasions can have severe direct and indirect impacts on natural communities and are one of the leading reasons behind the loss of biodiversity. We investigated the indirect effects of Argentine ant (Linepithema humile) invasions on coastal horned lizards (Phrynosoma coronatum) through an examination of prey selection in the field and laboratory. In non-invaded areas, coastal horned lizard diet consists predominately of ants (>94% by prey item in three reserves examined), particularly harvester ants in the genera Pogonomyrmex and Messor. In invaded areas, most native ants were displaced, and remaining horned lizards incorporated more non-ant arthropods and smaller ants into their diet. In addition, Argentine ants were never detected in horned lizard fecal pellets, suggesting that they are not a suitable replacement food source for the native species they are displacing. Analysis of prey selection in uninvaded areas reveals that ant species consumption by adult lizards is correlated with ant body size rather than ant abundance, foraging behavior, or defensive method. Differences in diet between adult and juvenile lizards suggest that a diversity of ant species is necessary to support both age classes. Laboratory experiments, consisting of paired presentations of four native ant species and the Argentine ant, confirmed patterns found in the field. Both adult and juvenile horned lizards preferred native ants to Argentine ants. Patterns of preference suggest that, in addition to L. humile's small size, other factors may contribute to the horned lizard's aversion to this species. Populations of coastal horned lizards have undergone severe declines in recent years. While habitat loss has been the primary concern, the invasion of Argentine ants into their remaining suitable habitat needs to be monitored. Reserves need to be managed to prevent the penetration of Argentine ants in order to minimize their direct and indirect effects on natural communities.

Simple SummaryWe aimed at assessing the impact of the Argentine ant invasion on the native ant community in citrus ecosystems. We compared the Argentine ant’s invaded and uninvaded citrus orchards in the south of Portugal, estimating species richness and the frequency of ant assemblages foraging on the tree canopy. The results suggest that the Argentine ant has a negative impact on the native ant community structure, markedly reducing the diversity and frequency of native species. This impact was more or less pronounced depending on the season. Possible implications for citrus pest management are discussed.The invasion of the Argentine ant, Linepithema humile (Mayr) (Hymenoptera, Formicidae) can alter the entire ecosystem with serious impacts on the native community structure (e.g., ant diversity) and processes (e.g., trophic interactions) leading to biodiversity loss and pest outbreaks. Most studies addressing these impacts have been conducted in natural or semi-natural areas, few are those conducted in agricultural ecosystems, such as citrus orchards. These are dominant agricultural ecosystems in Mediterranean landscapes. Furthermore, most studies have been conducted in a short span, not evidencing seasonal fluctuations. In this work, we assessed the ecological impact of the Argentine ant on the native ant communities in citrus orchards, in the region of Algarve, southern Portugal. By using principal response curve, we compared seasonal variation on ant assemblages in invaded and uninvaded citrus orchards foraging on tree canopy from a two-year sampling. The Argentine ant had a marked negative impact on the native ant community foraging on citrus canopy. In the uninvaded orchards, the native ant community had a rich assemblage composed of 16 ant species, in its majority (72%) controlled by the dominant species Lasius grandis Forel, Tapinoma nigerrimum (Nylander) and/or Pheidole pallidula (Nylander). In the invaded orchards, the native ant community was poorer and highly modified, mostly dominated by the Argentine ant (80%). Apparently, the only native ant species not affected by the presence of the Argentine ant was Plagiolepis pygmaea (Latreille). A significant negative effect was found between the proportion of infested trees by L. humile and the number of native ant species per orchard. Differences in the native ant community in the invaded and uninvaded orchards persisted over seasons and years. However, negative impacts were higher in the spring and summer, and less pronounced in the autumn. We discuss implications for citrus pest management.

Biological invasions can have severe direct and indirect impacts on natural communities and are one of the leading reasons behind the loss of biodiversity. We inves- tigated the indirect effects of Argentine ant (Linepithema humile) invasions on coastal horned lizards (Phrynosoma coronatum) through an examination of prey selection in the field and laboratory. In non-invaded areas, coastal horned lizard diet consists predominately of ants (>94% by prey item in three reserves examined), particularly harvester ants in the genera Pogonomyrmex and Messor. In invaded areas, most native ants were displaced, and remaining horned lizards incorporated more non-ant arthropods and smaller ants into their diet. In addition, Argentine ants were never detected in horned lizard fecal pellets, sug- gesting that they are not a suitable replacement food source for the native species they are displacing. Analysis of prey selection in uninvaded areas reveals that ant species con- sumption by adult lizards is correlated with ant body size rather than ant abundance, foraging behavior, or defensive method. Differences in diet between adult and juvenile lizards suggest that a diversity of ant species is necessary to support both age classes. Laboratory exper- iments, consisting of paired presentations of four native ant species and the Argentine ant, confirmed patterns found in the field. Both adult and juvenile horned lizards preferred native ants to Argentine ants. Patterns of preference suggest that, in addition to L. humile's small size, other factors may contribute to the horned lizard's aversion to this species. Populations of coastal horned lizards have undergone severe declines in recent years. While habitat loss has been the primary concern, the invasion of Argentine ants into their remaining suitable habitat needs to be monitored. Reserves need to be managed to prevent the penetration of Argentine ants in order to minimize their direct and indirect effects on natural communities.

Spatial and temporal variation in the Argentine ant edge effect: Implications for the mechanism of edge limitation

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.