Pre-adaptive shift of a native predator (Araneae, Zodariidae) to an abundant invasive ant species (Hymenoptera, Formicidae)

Invasive species often displace native species and can affect ecological processes in invaded habitats. If invasive species become abundant, changes in prey availability may be particularly harmful to specialist predators. The Argentine ant, Linepithema humile Mayr, is an important invasive species on nearly all continents. Spiders of the genus Zodarion are specialised ant-eating predators native to the Mediterranean yet it is unknown if they can exploit invasive ant species. Here we studied spatial and temporal abundance of this invasive ant and the native spider, Zodarion cesari Pekár, during 4?years in four citrus groves. Circadian activity of both spiders and ants, and capture efficiency and prey specificity of the predator were also evaluated. The abundance of Z. cesari was strongly correlated to L. humile abundance. The predatory activity of spiders varied seasonally with differences on the relative frequency of spiders capturing ants depending on the time of the year. In laboratory, Z. cesari displayed most efficient capture upon the native ant Tapinoma nigerrimum (Nylander) and the invasive ant L. humile in comparison with five other native ant species. These results demonstrate that the native spider Z. cesari is successfully exploiting the invasive ant species L. humile and is likely a locally monophagous predator. We suggest that Z. cesari shifted away from native T. nigerrimum post invasion as both ant species are phylogenetically related and of similar size.     

Diversity and structure of ant communities associated with oak,pine, eucalyptus and arable habitats in Portugal

Summary In central Portugal, 28 species of ants were found in oak, 25 in pine, 35 in eucalyptus and 12 in arable habitats, a total of 43 species. They include five numerically dominant species, the exotic Argentine ant.Linepithema (Iridomyrmex) humile and four native species,Crematogaster scutellaris, Pheidole pallidula, Tapinoma nigerrimum andTetramorium hispanicum. L. humile occurred in 34% of sampled sites. When present, it was abundant in all three arboreal habitats, where it was associated with strikingly decreased ant species richness and equitability. Such diversity was greater inL. humile-colonised arable habitats, where the ant was much less abundant than in the arboreal habitats.WhereL. humile was absent,C. scutellaris andP. pallidula were usually co-dominants in oak and pine, andT. nigerrimum andT. hispanicum in the arable habitat, but all were absent or rare in eucalyptus. Ants were numerically least abundant in eucalyptus but, in the absence ofL. humile, species richness was greater and the community more equitable than in the other habitats.L. humile-absent arable habitats supported the poorest and least equitable ant community.     

Quantitative analysis of the effects of the exotic Argentine ant on seed-dispersal mutualisms

Although it is increasingly clear that exotic invasive species affect seed-dispersal mutualisms, a synthetic examination of the effect of exotic invasive species on seed-dispersal mutualisms is lacking. Here, we review the impacts of the invasive Argentine ant (Linepithema humile) on seed dispersal. We found that sites with L. humile had 92 per cent fewer native ant seed dispersers than did sites where L. humile was absent. In addition, L. humile did not replace native seed dispersers, as rates of seed removal and seedling establishment were all lower in the presence of L. humile than in its absence. We conclude that potential shifts in plant diversity and concomitant changes in ecosystem function may be a consequence of Argentine ant invasions, as well as invasions by other ant species. Because very few studies have examined the effects of non-ant invasive species on seed-dispersal mutualisms, the prevalence of disruption of seed-dispersal mutualisms by invasive species is unclear.     

Patterns of floral resource use by two dominant ant species in a biodiversity hotspot

Ecological dominance in ants is often fuelled by carbohydrate intake. Most studies have focused on the importance of invasive ant mutualistic associations with trophobionts whereas few studies have investigated the importance of floral nectar on invasion success. In this study, utilisation of temporarily available floral nectar by the invasive Argentine ant, Linepithema humile, was compared to that of the dominant native ant, Anoplolepis custodiens, within the Cape Floristic Region (CFR), a biodiversity hotspot. The effect of these two focal ant species on species composition and abundance of ground foraging ants as well as floral arthropod visitors in inflorescences of Proteacea species was assessed. Foraging activity, and trophic ecology inferred from the abundance of natural stable isotopes of Carbon (δ¹³C) and Nitrogen (δ¹⁵N), and the ratio of Carbon to Nitrogen (C:N) were compared between the two ant species during three flowering periods. Linepithema humile significantly reduced the abundance and species diversity of both above-ground and floral arthropod species abundance and composition. Linepithema humile increased its foraging activity with increasing nectar availability, switching its diet to a more herbivorous one. Anoplolepis custodiens did not respond as effectively to increasing floral nectar or negatively impact floral arthropod visitors. This study showed that the availability of floral nectar and ability of L. humile to more effectively utilise this temporarily available resource than native ants, can contribute significantly to the further spread and persistence of L. humile in natural environments in the CFR.     

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.     

Propagule Pressure and Climate Contribute to the Displacement of Linepithema humile by Pachycondyla chinensis

Identifying mechanisms governing the establishment and spread of invasive species is a fundamental challenge in invasion biology. Because species invasions are frequently observed only after the species presents an environmental threat, research identifying the contributing agents to dispersal and subsequent spread are confined to retrograde observations. Here, we use a combination of seasonal surveys and experimental approaches to test the relative importance of behavioral and abiotic factors in determining the local co-occurrence of two invasive ant species, the established Argentine ant (Linepithema humile Mayr) and the newly invasive Asian needle ant (Pachycondyla chinensis Emery). We show that the broader climatic envelope of P. chinensis enables it to establish earlier in the year than L. humile. We also demonstrate that increased P. chinensis propagule pressure during periods of L. humile scarcity contributes to successful P. chinensis early season establishment. Furthermore, we show that, although L. humile is the numerically superior and behaviorally dominant species at baits, P. chinensis is currently displacing L. humile across the invaded landscape. By identifying the features promoting the displacement of one invasive ant by another we can better understand both early determinants in the invasion process and factors limiting colony expansion and survival.     

The Ants of the Juan Fernández Islands: Genesis of an Invasive Fauna

A first study of the ants on the three main islands of the Juan Fernández Archipelago (Chile) has revealed the presence of just three species, Hypoponera confinis, Linepithema humile, and Tetramorium bicarinatum, all widely distributed invasive species. Most notable among them is the Argentine ant, L. humile, an ecologically destructive species that has established several populations and appears to be expanding.     

Resource competition assays between the African big‐headed ant,Pheidole megacephala (Fabricius) and the invasive Argentine ant,Linepithema humile (Mayr): mechanisms of inter‐specific displacement

1. The spread of Argentine ants, Linepithema humile (Mayr), in introduced areas is mainly through the displacement of native ant species owing to high inter‐specific competition. In South Africa, L. humile has not established in the climatically suitable eastern and northern escarpments dominated by the African big headed ant, Pheidole megacephala (Fabricius), probably owing to local biotic resistance. 2. Inter‐specific aggression, at the individual and colony level, and competition for a shared resource were evaluated in the laboratory. 3. Aggression between the two ant species was very high in all of the assays. Both species suffered similar mortality rates during one‐on‐one aggression assays, however, during symmetrical group confrontations, L. humile workers showed significantly higher mortality rates than P. megacephala workers. During asymmetrical group confrontations both species killed more of the other ant species when they had numeric advantage. Both ant species located the shared resource at the same time; however, once P. megacephala discovered the bait, they displaced L. humile from the bait through high inter‐specific aggression, thereafter dominating the bait for the remainder of the trial. 4. The results demonstrate the potential of P. megacephala to prevent the establishment and survival of incipient L. humile colonies through enhanced resource competition and high inter‐specific aggression. This is the first study to indicate potential biotic resistance to the spread of L. humile in South Africa.     

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).     

Natural biological control of the pine processionary moth Thaumetopoea pityocampa (Den. & Schiff.) by the Argentine ant Linepithema humile (Mayr) in Portugal

1 Defoliation by larvae of the pine processionary moth, Thaumetopoea pityocampa, is negligible in stands of Pinus pinaster colonized by the Argentine ant Linepithema humile which preys fiercely on the young moth larvae. In contrast, such damage is widespread where pine plantations are colonized by native ants, predominantly Tapinoma nigerrimum and Lasius niger, which seemingly disregard the larvae. 2 Where L. humile- and native ant-occupied sectors adjoin, there is a 20–50 m overlap in the transition area between L. humile- and native ant-occupied pines. This was most evident in a > 500 ha plantation where there was severe or very severe T. pityocampa attack in native ant sectors contrasting with none in adjoining L. humile sectors. 3 Predation by L. humile is no doubt enhanced by its existence as super-colonies over very large areas, by its foraging activity and recruitment on trees throughout the time when T. pityocampa and other prey are present, and by honeydew- producing Homoptera which help retain foraging L. humile workers in pine tree crowns. 4 The role of L. humile could be enhanced by cultivations that disturb the soil and restrain ground vegetation.     

Interaction between a threatened endemic plant (<Emphasis Type="Italic">Anchusa crispa</Emphasis>) and the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>)

Seed dispersal mutualisms are essential to ensure the survival of diverse plant species and communities worldwide. Here, we investigated whether the invasive Argentine ant can replace native ants by fulfilling their functional role in the seed dispersal of the rare and threatened endemic myrmecochorous plant, Anchusa crispa, in Corsica (France). Our study addressed the potential of Linepithema humile to disperse elaiosome-bearing seeds of A. crispa, examining L. humile’s effects on (1) the composition of communities of ants removing seeds, (2) the number of seed removals, (3) seed preference, (4) the distance of seed dispersion, and (5) seed germination. We caught seven native species at the control site, but only the Argentine ant at invaded sites. L humile removed A. crispa seeds in greater numbers than did native ants, respectively 66 and 23%, probably due to their higher worker density. The invader was similar to native ants with respect to distance of seed transport. Finally, rates of seed germination were not significantly different between seeds previously in contact with either Argentine ants or not. Taken all together, these results suggest that the Argentine ant is unlikely to pose a threat to A. crispa population. These results have important implications for the management of this rare and threatened endemic plant and provide an example of non-negative interactions between invasive and native species.     

Floral visitation by the Argentine ant reduces pollinator visitation and seed set in the coast barrel cactus, Ferocactus viridescens

Mounting evidence indicates that trade-offs between plant defense and reproduction arise not only from resource allocation but also from interactions among mutualists. Indirect costs of plant defense by ants, for example, can outweigh benefits if ants deter pollinators. Plants can dissuade ants from occupying flowers, but such arrangements may break down when novel ant partners infiltrate mutualisms. Here, we examine how floral visitation by ants affects pollination services when the invasive Argentine ant (Linepithema humile) replaces a native ant species in a food-for-protection mutualism with the coast barrel cactus (Ferocactus viridescens), which, like certain other barrel cacti, produces extrafloral nectar. We compared the effects of floral visitation by the Argentine ant with those of the most prevalent native ant species (Crematogaster californica). Compared to C. californica, the Argentine ant was present in higher numbers in flowers. Cactus bees (Diadasia spp.), the key pollinators in this system, spent less time in flowers when cacti were occupied by the Argentine ant compared to when cacti were occupied by C. californica. Presumably as a consequence of decreased duration of floral visits by Diadasia, cacti occupied by L. humile set fewer seeds per fruit and produced fewer seeds overall compared to cacti occupied by C. californica. These data illustrate the importance of mutualist identity in cases where plants balance multiple mutualisms. Moreover, as habitats become increasingly infiltrated by introduced species, the loss of native mutualists and their replacement by non-native species may alter the shape of trade-offs between plant defense and reproduction.     

Carbohydrate supply limits invasion of natural communities by Argentine ants

The ability of species to invade new habitats is often limited by various biotic and physical factors or interactions between the two. Invasive ants, frequently associated with human activities, flourish in disturbed urban and agricultural environments. However, their ability to invade and establish in natural habitats is more variable. This is particularly so for the invasive Argentine ant (Linepithema humile). While biotic resistance and low soil moisture limits their invasion of natural habitats in some instances, the effect of food availability has been poorly explored. We conducted field experiments to determine if resource availability limits the spread and persistence of Argentine ants in remnant natural forest in North Carolina. Replicated transects paired with and without sucrose solution feeding stations were run from invaded urban edges into forest remnants and compared over time using baits and direct counts at feeding stations. Repeated under different timing regimes in 2006 and 2007, access to sucrose increased local Argentine ant abundances (1.6–2.5 fold) and facilitated their progression into the forest up to 73 ± 21% of 50-m transects. Resource removal caused an expected decrease in Argentine ant densities in 2006, in conjunction with their retreat to the urban/forest boundary. However, in 2007, Argentine ant numbers unexpectedly continued to increase in the absence of sugar stations, possibly through access to alternative resources or conditions not available the previous year such as honeydew-excreting Hemiptera. Our results showed that supplementing carbohydrate supply facilitates invasion of natural habitat by Argentine ants. This is particularly evident where Argentine ants continued to thrive following sugar station removal.     

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.     

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.     

Colony structure in introduced and native populations of the invasive Argentine ant, <Emphasis Type="Italic">Linepithema humile</Emphasis>

Summary. The Argentine ant, Linepithema humile, severely decreases the abundance and diversity of native ant fauna in areas where it invades, but coexists with a more diverse assemblage of ants in its native range. The greater ecological dominance of L. humile in the introduced range may be associated with differences in colony structure and population density in the introduced range relative to the native range. In this study, I compared aspects of L. humiles colony structure, including density, the spatial pattern of nests and trails, and patterns of intraspecific aggression in parts of the introduced and native ranges. I also compared the number of ant species coexisting with L. humile. Introduced and native populations did not differ significantly in nest density, ant density, nest size, and nearest-neighbor distances. In three of the four study populations in the native range and all of the study populations in the introduced range, colonies were organized into supercolonies: they consisted of multiple, interconnected nests that were dense and spatially clumped, and aggression among conspecifics was rare. In one population in the native range, colonies were organized differently: they occupied single nest sites, nests were sparse and randomly dispersed, and ants from neighboring nests were aggressive toward each other. Species richness was significantly higher in the native range than in the introduced range, even in areas where L. humile formed dense supercolonies. The results suggest that differences in species coexistence between ranges may due to factors other than L. humiles colony structure. One likely factor is the superior competitive ability of other ant species in the native range.Received 23 January 2004; revised 30 March 2004; accepted 20 April 2004.     

Application of the LAMP assay for the detection of the Argentine ant, <Emphasis Type="Italic">Linepithema humile</Emphasis> (Hymenoptera: Formicidae), from captures of pan traps

We applied the loop-mediated isothermal amplification (LAMP) assay to monitor invasions of Linepithema humile (Mayr), the Argentine ant, a notorious invasive insect worldwide. Species-specific LAMP primers were designed on the basis of the partial sequence of the cytochrome c oxidase subunit I region of L. humile. The species specificity and sensitivity of these primers were determined in the laboratory and considered adequate for practical use. We also confirmed that the assay successfully detected L. humile from captures of pan traps, which contained L. humile and several non-target ant species. The assay detected the target species even when the captures contained only a leg or an antenna. Since the LAMP assay is simple and rapid, this assay will contribute to the early detection and accurate identification of L. humile.     

Invasive ants carry novel viruses in their new range and form reservoirs for a honeybee pathogen

When exotic animal species invade new environments they also bring an often unknown microbial diversity, including pathogens. We describe a novel and widely distributed virus in one of the most globally widespread, abundant and damaging invasive ants (Argentine ants, Linepithema humile). The Linepithema humile virus 1 is a dicistrovirus, a viral family including species known to cause widespread arthropod disease. It was detected in samples from Argentina, Australia and New Zealand. Argentine ants in New Zealand were also infected with a strain of Deformed wing virus common to local hymenopteran species, which is a major pathogen widely associated with honeybee mortality. Evidence for active replication of viral RNA was apparent for both viruses. Our results suggest co-introduction and exchange of pathogens within local hymenopteran communities. These viral species may contribute to the collapse of Argentine ant populations and offer new options for the control of a globally widespread invader.     

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.     

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.