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.     

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.     

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.     

The diminutive supercolony: the Argentine ants of the southeastern United States

Native to Argentina and Brazil, the Argentine ant (Linepithema humile) is an invasive species that has become established on six continents and many oceanic islands. In several parts of its introduced range, including the western United States, southern Europe and Chile, the Argentine ant is unicolonial, forming extensive supercolonies. We examined population genetic structure and intercolony aggression in two regions of the introduced range of this species in the United States: California and the southeastern United States. Our results show that the southeastern L. humile population has high genotypic variability and strong intercolony aggression relative to the California population. In the California population, intercolony aggression was absent and 23 alleles were found across seven polymorphic microsatellite loci. However, in the Southeast, aggression between colonies was high and 47 alleles were present across the same seven loci in an equal number of colonies. We suggest that distinctly different colonization patterns for California and the Southeast may be responsible for the striking disparity in the genetic diversity of introduced populations. Southeastern colonies may have descended from multiple, independent introductions from the native range, undergoing a bottleneck at each introduction. In contrast, the California supercolony may have originated from one or more colonies inhabiting the southeastern United States, thus experiencing a double bottleneck. The differences in present-day distribution patterns between California and the Southeast may be due to the combined effect of two factors: lower winter temperatures in the Southeast and/or competition with another successful and widely distributed ant invader, the fire ant Solenopsis invicta.     

Argentine ant foraging activity and interspecific competition in complete vs. queenless and broodless colonies

Introduction experiments may prove useful in understanding the mechanisms underlying the successful establishment of invasive ant species into new areas. These manipulative introductions could be particularly helpful in exploring the interactions between invasive species and the local fauna and flora. However, the inherent risk of accidental establishment in such experiments poses unacceptable ethical concerns. Some of the worst invasive species are tramp ant species, which can adversely affect biodiversity and community structure after establishment. We conducted laboratory and field experiments investigating a safe methodology for carrying out introduction experiments using the sterile workers of the invasive Argentine ant, Linepithema humile, as a model. We found no difference in foraging rate between worker-only colonies of L. humile and complete colonies, containing queens, workers and brood. Worker-only L. humile colonies showed the same exploitative and interference ability as complete colonies in bait dominance trials with the odorous house ant, Tapinoma sessile, in both laboratory and field trials. We suggest that for those invasive ant species with sterile workers, worker-only colonies may be substituted for complete colonies in short-term field experiments in new areas. Received 18 January 2007; revised 19 June 2007; accepted 22 June 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     

江苏省外来种子植物的初步调查和分析

对江苏省外来种子植物的种类、来源地及分布状况进行了初步调查统计。调查结果显示,江苏省共有外来种子植物393种,大多数来源于欧洲和美洲,主要通过有意引种和无意引入2种方式引入。有意引种的栽培植物有276种,其中观赏植物132种、蔬菜46种、林木35种、工业原料植物17种、药用植物12种、牧草植物11种、粮食作物11种、果树7种以及油料作物5种。另外,外来种子植物中有逃逸种62种、归化种22种、入侵种33种;81．8％的入侵种在江苏全省均有分布;入侵种以菊科（Compositae）和禾本科（Poaceae）植物为主,分别有11种和7种。在此调查结果基础上,对江苏省外来种子植物的分类和危害进行了讨论,并对江苏省外来物种的利用和管理提出了一些建议。     

Lack of inbreeding avoidance in the Argentine ant Linepithema humile

Although workers might increase their inclusive fitness by favoringcloser over more distant kin, evidence suggest that nepotismgenerally does not occur within colonies of social insects.It has been suggested that this may be due to the cost of recognitionerrors. We tested whether recognition occurs in a system wherea better than random ability to recognize kin should be selected for. Using DNA microsatellites, we show that sexuals of theArgentine ant Linepithema humile fail to use genetic cues toavoid sib-mating. When offspring of two queens were allowedto mate, the percentage of matings among siblings was not significantlylower than expected under the hypothesis of random mating.The finding that sexuals fail to use genetic cues to avoid sib-matings cannot be attributed to the cost of recognitionerrors because any recognition system that would lead to abetter than random ability to avoid sib-mating should be selectedfor when there are costs to inbreeding. These data are thusconsistent with the view that kin recognition mediated solelyby genetic cues might be intrinsically error prone within coloniesof social insects.     

The apparent establishment and subsequent eradication of the Australian giant bulldog ant Myrmecia brevinoda Forel (Hymenoptera: Formicidae) in New Zealand

Abstract

The Australian giant bulldog ant Myrmecia brevinoda Forel was first observed in New Zealand in 1940 (Keall 1981). Further observations of this species were made at a location in Devonport in 1948 and 1965, and a nest destroyed at this site in 1981. Since this time no further observations have been made, yet the status of this species is commonly listed as “uncertain establishment” on New Zealand faunal lists. We describe the control procedure used in 1981. In addition, to determine its current status we resurveyed ants in the area of earlier infestation using visual searches, pitfall trapping, and a reward for discovery posted to local residents. No evidence of M. brevinoda was found, though several other invasive species were found, including the Argentine ant Linepithema humile (Mayr), Pheidole rugosula Forel, and a recently established Australian cryptic Solenopsis species. Given the results of our survey and the absence of other reports of M. brevinoda over the last 24 years, we consider this species to be extinct or eradicated from New Zealand.     

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.     

利用传统生物防治控制外来杂草的入侵

随着国际贸易的日益频繁,外来有害植物入侵,严重威胁我国的自然环境和生物多样性。利用从原产地引入食性较专一的天敌来控制外来杂草是杂草生物防治的主要方式之一,有保护环境一劳永逸的效果。简要介绍了国际生物防治概况,统计表明全世界至少有133种目标杂草进行生物防治,主要分布在菊科、仙人掌科和含羞草科,63科369种无脊椎动物和真菌作为杂草生物防治的天敌,利用最多的天敌是鞘翅目象甲科和叶甲科昆虫,其中大多数项目是治理外来杂草的。杂草生物防治最活跃的国家依次为美国、澳大利亚、南非、加拿大和新西兰。重点论述了利用传统生物防治方法防治外来杂草的经典项目、国内外研究概况,以及目前面临的问题和应用前景。我国杂草生物防治起步晚,传统杂草生防的目标杂草有4种,紫茎泽兰、空心莲子草、豚草和水葫芦,其中,空心莲子草的生物防治获得成功。共引进天敌14种,输出天敌23种,与世界上生物防治先进的国家比尚有距离。中国应充分借鉴国际成功经验,对外来杂草开展生物防治。中国的生物多样性在世界上占有十分独特的地位,将在生物多样性保护中发挥重要作用。     

Role of exotic tree stands on the current distribution and social behaviour of Swainson's hawk, Buteo swainsoni in the Argentine Pampas

Aim The Argentine Pampas was extensively and abruptly altered by European colonization. Between 1880 and 1885 the indigenous human inhabitants were completely displaced, and native grasslands were replaced by exotic pasture plants and crops. One of the most important ecological changes in the Pampas landscape, the introduction of tall exotic tree species, has received little attention, and its effect on wildlife has never been assessed. We have made an intensive survey of habitat use of Swainson's hawk, Buteo swainsoni in its most important non‐breeding quarters, the Pampas of Argentina, aiming to characterize the sites used by hawks for communal roosting. Location Pampas grasslands, Argentina. Methods We surveyed 30,000 km of roads by car during the austral summer from 2001 to 2004, covering the main non‐breeding area occupied by Swainson's hawks. Their roost sites were located by direct observation of birds roosting in tree stands close to the roads, by surveying potential roost sites around places where groups of pre‐ and post‐roosting hawks were recorded, and by gathering information from local farmers. Results Swainson's hawks exclusively used stands of exotic tree species for roosting. Eucalyptus viminalis was present in all 34 roosts surveyed, and in 59% of them it was the only species present. The remaining exotic tree species were the Siberian elm, Ulmus pumila, pines (Pinus spp.) and cypress (Cupressus spp.). Flock sizes at roost sites were unusually high for a raptor, with an average of 658 individuals (range 8–5000 hawks, n = 27 flocks). Main conclusions The introduction of exotic trees may have resulted in the expansion of the suitable habitat for Swainson's hawks, permitting a recent colonization of the Argentine Pampas. Tree stands may have also changed the communal roosting behaviour of this raptor, by virtue of their providing new structural elements in a region that almost completely lacked trees prior to European occupation.     

The worldwide transfer of ants: geographical distribution and ecological invasions

Aim This is the first comprehensive account of the biogeography of ants transferred and at least temporarily established outside their native habitat. Location Using museum and literature records, I established the distributions of transferred ant species. Methods I used taxonomic and functional groups to assess how geographical spread as a transferred species is affected by taxonomy and life history. Results 147 ant species in forty-nine genera have been recorded outside of their native habitat. The proportion of transferred ants is similar to the number of genera and species in each subfamily. The species-rich subfamily Myrmicinae contains nearly 50% of all transferred species, while many of the species-poor subfamilies have absolutely no transferred species. A disproportionate high number of transferred ants originate from the Neotropical and Oriental biogeographic regions. The Pacific Islands are the recipients of the most transferred ant species. Most transferred ants belong to the CRYPTIC, OPPORTUNIST, and GENERALIZED MYRMICINE functional groups, while there are no recorded transfers of army ants or leaf-cutting ants. Both invasive and human commensal ‘tramp’ ant species are nonrandom subsets of transferred ants. Main conclusions ‘Tramp’ species and invasive species tend to have widespread geographical distributions, and share life history characteristics including queen number, nest structure, and foraging behaviour. Combining observations of functional groups and biogeography may lead to a better understanding of the factors contributing to the spread of transferred species.     

Traits allowing some ant species to nest syntopically with the fire ant Solenopsis saevissima in its native range

Supercolonies of the red fire ant Solenopsis saevissima (Smith) develop in disturbed environments and likely alter the ant community in the native range of the species. For example, in French Guiana only 8 ant species were repeatedly noted as nesting in close vicinity to its mounds. Here, we verified if a shared set of biological, ecological, and behavioral traits might explain how these 8 species are able to nest in the presence of S. saevissima. We did not find this to be the case. We did find, however, that all of them are able to live in disturbed habitats. It is likely that over the course of evolution each of these species acquired the capacity to live syntopically with S. saevissima through its own set of traits, where colony size (4 species develop large colonies), cuticular compounds which do not trigger aggressiveness (6 species) and submissive behaviors (4 species) complement each other.     

Specialized predation on Wasmannia auropunctata by the army ant species Neivamyrmex compressinodis

Abstract:  We report here the first case of an efficient and specialized predator of the invasive ant species Wasmannia auropunctata : the army ant Neivamyrmex compressinodis . Our results are based on a study that we conducted in French Guiana, a part of the Wasmannia's native range. When N . compressinodis workers attacked W . auropunctata nests, the assaulted workers panicked and left the nests, some of them carrying brood. Nevertheless, during its raids on W . auropunctata , N . compressinodis was able to capture nearly all of the W . auropunctata brood and winged sexuals, whereas none of the attacks by N . compressinodis on other sympatric ant species were successful. Laboratory experiments revealed that the workers of eight compared sympatric species attacked the N . compressinodis individuals and that N . compressinodis workers accepted W . auropunctata brood as well as that of most of the tested species, showing that its specificity probably depends on the reaction of the W . auropunctata workers.     

Behavioral and Genetic Differentiation Between Native and Introduced Populations of the Argentine Ant

In this paper, we examine the hypothesis that reduced intraspecific aggression underlies the competitive prowess of Argentine ants in their introduced range. Specifically, we test three predictions of this hypothesis by comparing the genetic diversity, behavior, and ecology of Argentine ants in their native range to introduced populations. Differences between native and introduced populations of Argentine ants were consistent with our predictions. Introduced populations of the Argentine ant appear to have experienced a population bottleneck at the time of introduction, as evidenced by much reduced variation in polymorphic microsatellite DNA markers. Intraspecific aggression was rare in introduced populations but was common in native populations. Finally, in contrast to the Argentine ant's ecological dominance throughout its introduced range, it did not appear dominant in the native ant assemblages studied in Argentina. Together these results identify a possible mechanism for the widespread success of the Argentine ant in its introduced range.     

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.     

Intraspecific competition influences the symmetry and intensity of aggression in the Argentine ant

Cooperative social groups rely on the ability to distinguishmembers from nonmembers. Accordingly, social insects have evolveda variety of systems that allow discrimination of nest matesfrom non–nest mates. In this study, we show that experiencecan modify patterns of intraspecific aggression in Argentineants (Linepithema humile). In laboratory experiments, we foundthat aggression between colonies was often asymmetrical, butin all five cases, this asymmetry shifted to symmetrical aggressionafter contact with a hostile colony. Moreover, in the field,aggression between workers collected from colony borders wassymmetrical, whereas polarized aggression occurred between workerscollected 500 m away from colony borders. Coinciding with thisshift in aggression symmetry, we also observed an increase inboth the overall level of aggression and the frequency of aggressionin both the field and laboratory bioassays. We found littleevidence for colony-level competitive asymmetries stemming frompolarities in aggression at the worker level, either in thelaboratory or in the field. These results illustrate that recognitionsystems in Argentine ants are surprisingly dynamic and provideexperimental evidence for how recognition can be adjusted inresponse to specific circumstances—in this case the presenceof intraspecific competitors.