Importance of large colony size for successful invasion by Argentine ants (Hymenoptera: Formicidae): Evidence for biotic resistance by native ants

Abstract The Argentine ant, Linepithema humile (Mayr), is a widespread invasive ant species that has been associated with losses of native ant species and other invertebrates from its introduced range. To date, various abiotic conditions have been associated with limitations to the spread of Argentine ants, however, competitive interactions with native ant fauna may also affect the spread of Argentine ants. Here, we experimentally manipulated colony sizes of Argentine ants in the laboratory to assess whether Argentine ants were able to survive and compete for resources with a widespread, dominant native ant, Iridomyrmex‘rufoniger’. The results showed that over 24 h, the proportions of Argentine ants that were alive, at baits, and at sugar water decreased significantly in the presence of Iridomyrmex. In addition, Argentine ant mortality increased over time, however, the proportion of the colony that was dead decreased with the largest colony size. Argentine ants were only able to overcome Iridomyrmex when their colony sizes were 5–10 times greater than those of the native ants. We also conducted trials in which colonies of Argentine ants of varying sizes were introduced to artificial baits occupied by Iridomyrmex in the field. The results showed that larger Argentine ant colonies significantly affected the foraging success of Iridomyrmex after the initial introduction (5 min). However, over the first 20 min, when the Argentine ants were present at the baits, and over the entire 50 min experimental period, the numbers of Iridomyrmex at baits did not differ significantly with the size of the Argentine ant colony. This is the first experimental study to investigate the role of colony size in the invasion biology of Argentine ants in Australia, and the results suggest that Iridomyrmex may reduce the spread of Argentine ants, and that Argentine ants may need to attain large colony sizes in order to survive in the presence of Iridomyrmex. We address the implications of these findings for the invasion success of Argentine ants in Australia, and discuss the ability of Argentine ants to attain large colony sizes in introduced areas.     

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.     

Impacts of Argentine ants on mealybugs and their natural enemies in California's coastal vineyards

Abstract 1. The Argentine ant, Linepithema humile, tends honeydew‐excreting homopterans and can disrupt the activity of their natural enemies. This mutualism is often cited for increases in homopteran densities; however, the ant’s impact on natural enemies may be only one of several effects of ant tending that alters insect densities. To test for the variable impacts of ants, mealybug and natural enemy densities were monitored on ant‐tended and ant‐excluded vines in two California vineyard regions. 2. Ant tending increased densities of the obscure mealybug, Pseudococcus viburni, and lowered densities of its encyrtid parasitoids Pseudaphycus flavidulus and Leptomastix epona. Differences in parasitoid recovery rates suggest that P. flavidulus was better able to forage on ant‐tended vines than L. epona. 3. Densities of a coccinellid predator, Cryptolaemus montrouzieri, were higher on ant‐tended vines, where there were more mealybugs. Together with behavioural observations, the results showed that this predator can forage in patches of ant‐tended mealybugs, and that it effectively mimics mealybugs to avoid disturbance by ants. 4. Ant tending increased densities of the grape mealybug, Pseudococcus maritimus, by increasing the number of surviving first‐instar mealybugs. Parasitoids were nearly absent from the vineyard infested with P. maritimus. Therefore, ants improved either mealybug habitat or fitness. 5. There was no difference in mealybug distribution or seasonal development patterns on ant‐tended and ant‐excluded vines, indicating that ants did not move mealybugs to better feeding locations or create a spatial refuge from natural enemies. 6. Results showed that while Argentine ants were clearly associated with increased mealybug densities, it is not a simple matter of disrupting natural enemies. Instead, ant tending includes benefits independent of the effect on natural enemies. Moreover, the effects on different natural enemy species varied, as some species thrive in the presence of ants.     

Relative risk of invasive ants (Hymenoptera: Formicidae) establishing in New Zealand

Abstract

Human‐assisted spread of species poses a major challenge to border security agencies. Ideally, limited resources need to be targeted at species posing the most risk. Climate matching is an important component of assessing risk but often little or no biological information is available to enable detailed modelling. To assess if distribution records alone provide useful establishment predictions we compare the climate in New Zealand and its outlying islands to that found in the current native and introduced ranges of 12 tramp ant species, three of which are already established in New Zealand, using the climate module of BIOSECURE, a risk assessment tool. Eleven species showed a similar general pattern, with mean annual temperature being the climatic variable with the least overlap between the estimated realised niche and the predicted New Zealand niche. The twelfth species, of temperate origin, is less restricted by temperature, but much of New Zealand may have too high a rainfall. The present and future threats posed by tramp ants are discussed in relation to climate limitations.     

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.     

Fecundity and longevity of Argentine ant (Hymenoptera: Formicidae) queens in response to irradiation

Irradiation is a post‐harvest quarantine treatment option to control ants and other hitchhiker pests on fresh horticultural products traded between countries. As little is known about irradiation effects on ants, radiotolerance of the Argentine ant, Linepithema humile (Mayr) (Hymenoptera: Formicidae: Dolichoderinae), was studied to determine a dose sufficient for its control. Queens collected from Buenos Aires, Argentina, were irradiated with 30, 60, 90 Gy or left untreated as controls, and then followed for 8 weeks to evaluate their survival and fecundity. Overall queen survival and brood viability decreased with increasing irradiation dose. The number of eggs was reduced by 50%, 69% and 56% in the 30, 60 and 90 Gy doses, respectively, compared with untreated control queens. The percentage of eggs that developed into larvae decreased from 41.1% in the control to 22.5%, 1.4%, and 0% in the 30, 60, and 90 Gy treatments, respectively. Thus, the number of larvae was reduced by 69% in the 30 Gy treatment compared with the control, only one larva was observed in the 60 Gy treatment, and none in the 90 Gy treatment. Only one pupa was observed in the 30 Gy treatment and none in the 60 and 90 Gy treatments during the 8‐week experiment. Queens irradiated with 60 and 90 Gy had significantly reduced longevity compared with queens treated with lower doses or untreated queens. Radiation dose ≥90 Gy stopped brood development in Argentine ant queens and should be sufficient as a phytosanitary treatment. The radiotolerance of Argentine ant appears to be similar to that of two other important invasive ants.     

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.     

Effect of a highway on ant (Hymenoptera: Formicidae) species composition and abundance, with a recommendation for roadside verge width

Species composition and relative abundances of ground-dwelling ants at various distances from a major urban highway in South Africa were sampled using pitfall trapping. Diversity, richness and evenness indices indicated differences between ant assemblages, with samples near to the highway (<4 m) having the highest species diversity, and farthest ones (32 m) having the lowest. As measured environmental variables did not explain much of the variation in the species spatial pattern, it appears that the ants were influenced mostly by interspecific competition and/or distribution of food resources, especially road kills. Pheidole megacephala was highly dominant at all sites and was negatively associated with several other species. This and other dominant species appeared to be influencing the abundance of the rarer ones. Dominance and high abundances of species in the genera Pheidole, Monomorium, Tetramorium and Paratrechina, along with the occurrence of certain grasses, indicated that the whole sampling area was disturbed. In a suburban setting, a narrow roadside verge (<4m) is probably adequate. However, for maximum diversity in rural areas, roadsides should be as wide as possible and also rich and heterogenous in native vegetation.     

Introduced ants (Hymenoptera: Formicidae) of mainland France and Belgium,with a focus on greenhouses

The monitoring of introduced species is becoming more important as global trade intensifies. Although ants make up a larger proportion of species on the list of the most invasive species in the world compared with other groups, little is known about the occurrence of those introduced in France, especially inside heated buildings. Here we review the literature available for mainland France and Belgium and report the results of a survey conducted with the help of tropical building managers between 2014 and 2016. We report for the first time in France the presence of Technomyrmex vitiensis and Plagiolepis alluaudi in multiple greenhouses. Technomyrmex difficilis was also found in one greenhouse for the first time in Europe. The diversity of introduced ants in greenhouses is very low, and these buildings are most often dominated by one or two species. We compared the most recent data and those collected throughout the twentieth century and showed that ant communities have changed substantially. Greenhouses could be responsible for the introduction of invasive species because they regularly import exotic plants, but we found no evidence that the three species of invasive ants present outdoors in France were introduced from greenhouses, where they rarely occur. We also report that introduced ants are pests in greenhouses because they disperse scale insects and kill biological control agents. The suppression of these ants could ease the maintenance of plants inside greenhouses.     

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.     

Spatial distribution of ants (Hymenoptera: Formicidae), vine mealybugs and mealybug parasitoids in vineyards

The mutualistic association between some ant species and honeydew‐producing Hemiptera has been shown to influence the distribution patterns and abundance of these hemipterans and their natural enemies. We studied the spatial distribution patterns of three ant species, mealybugs and mealybug parasitoids for two consecutive growing seasons on three wine grape farms in the Western Cape, South Africa. During the study period, no ant or mealybug controls were applied. Ant and mealybug monitoring was conducted on a total of 21 ha using a presence/absence sampling system, while parasitoids were collected from infested mealybug females. Spatial analysis by distance indices was used to analyse spatial distribution of insects and ArcView? was used to map the gap, patch and local association indices where significant association and disassociation occurred. Significant associations were found between some ants and parasitoids, while significant disassociations between the ants Crematogaster peringueyi and Linepithema humile; and also between Crematogaster peringueyi and Anoplolepis steingroeveri were found. Interspecific competition between ant species could play a role in the distribution of parasitoids and mealybugs. Our results stress the importance of monitoring for ants and mealybugs and further highlight the importance of restricted chemical applications against ants during the growing season.     

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.     

Effects of Argentine ants (Hymenoptera: Formicidae) on myrmecophilous lycaenid butterfly,Narathura bazalus (Lepidoptera: Lycaenidae), in western Japan

Effects of the invasive Argentine ant, Linepithema humile (Mayr), on a myrmecophilous butterfly, Narathura bazalus (Hewiston), were investigated in the field in western Japan. Larvae of N. bazalus were attended by workers of Argentine ants in invaded parks and of ten native and one cosmopolitan ant species in uninvaded parks. The abundance of eggs and larvae were not significantly different between invaded and uninvaded parks. Pupal weight and parasitized ratio by tachinid flies were also not different between the two types of parks. These results indicate that the role of Argentine ants for the butterfly might be almost equivalent to the native ants.     

Pitfall trapping for ants (Hymenoptera,Formicidae) in mesic Australia: the influence of trap diameter

Catches of ants in the two most commonly-used forms of pitfall trap (‘test-tube’ traps, ca 18 mm diameter; ‘coffee cup’ traps, ca. 70 mm diameter) are compared from samples in open grassy woodland in southern Victoria, Australia. The 25 morphospecies found in the narrower traps were all represented among the 31 morphospecies collected in the larger traps. Either pattern is adequate to collect samples for broad inter-treatment comparisons and documentation of the more typical and representative fauna, but larger traps may have some advantage if more complete inventory is sought.     

Determinants of local ant (Hymenoptera: Formicidae) species richness and activity density across Europe

1. Species richness is influenced by local habitat features and large‐scale climatic gradients. Usually, both influences are studied in isolation because of the divergent spatial scales at which they occur. Here, we compared the influence of large‐scale climate and local habitat type on European ants using a continent‐wide, standardised sampling programme. 2. We investigated species richness and activity density from pitfall traps distributed over four habitat types at 17 locations from northern Sweden to Spain and Greece. Species richness and activity density were analysed with respect to ambient energy [equilibrium evapotranspiration (EET)] and productive energy (net primary productivity). Furthermore, we compared ant richness and activity density between the four habitat types: arable land, scrubland, grassland, and forest. 3. Species richness and activity density of ants increased with equilibrium evapotranspiration (EET), explaining 30.2% of the total variation in species richness and 24.2% of activity density. Habitat type explained an additional 19.2% of the variation in species richness and 20.2% of activity density, and was not related to productivity. Species richness and activity density were highest in scrubland and significantly lower in forest and (marginally significant) in arable land. 4. The increase in EET and the decrease in forest confirms the pronounced thermophily of ants, whereas the decrease in arable land is probably caused by soil disturbance.     

Aphis clerodendri Matsumura (Hemiptera: Aphididae), attendant ants (Hymenoptera: Formicidae) and associates on Clerodendrum (Verbenaceae) in Australia

Abstract Aphis clerodendri Matsumura is newly recorded from Australia and is known from the Northern Territory, on islands in Torres Strait, and in rainforest in northern Queensland and New South Wales. It induces the formation of leaf pseudogalls on native species of Clerodendrum and is commonly attended by ants, which penetrate and may polydomously nest in the galls. Previously known only from eastern Asia, A. clerodendri can now be classified as native to Australia and Australasian in natural distribution. The species is also newly recorded from Papua New Guinea and Vietnam.     

Competition between grass‐cutting Atta vollenweideri ants (Hymenoptera: Formicidae) and domestic cattle (Artiodactyla: Bovidae) in Argentine rangelands

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Immediate impacts of invasion by Wasmannia auropunctata (Hymenoptera: Formicidae) on native litter ant fauna in a New Caledonian rainforest

Abstract For the last 30 years, Wasmannia auropunctata (the little fire ant) has spread throughout the Pacific and represents a severe threat to fragile island habitats. This invader has often been described as a disturbance specialist. Here we present data on its spread in a dense native rainforest in New Caledonia. We monitored by pitfall trapping the litter ant fauna along an invasive gradient from the edge to the inner forest in July 1999 and March 2000. When W. auropunctata was present, the abundance and richness of native ants drops dramatically. In invaded plots, W. auropunctata represented more than 92% of all trapped ant fauna. Among the 23 native species described, only four cryptic species survived. Wasmannia auropunctata appears to be a highly competitive ant that dominates the litter by eliminating native ants. Mechanisms involved in this invasive success may include predation as well as competitive interactions (exploitation and interference). The invasive success of W. auropunctata is similar to that of other tramp ants and reinforces the idea of common evolutionary traits leading to higher competitiveness in a new environment.     

Impact of Oreochromis niloticus (Linnaeus, 1758) (Pisces: Cichlidae) invasion on taxonomic and functional diversity of native fish species in the upper Kabompo River,northwest of Zambia

Invasive alien species have been revealed to drastically alter the structure of native communities; however, there is scarce information on whether taxonomic and functional spaces occupied by native species are equally filled by exotic species. We investigated the diversity of native species to understand the impact of exotic Oreochromis niloticus in the upper Kabompo River, northwest of Zambia using taxonomic and functional diversity indices. To achieve this, two tests were performed (Test 1, compared natives in invaded and uninvaded sections; Test 2, compared natives in invaded section). A total of 17 species were collected for functional diversity computation, out of which fourteen (14) functional trait measurements linked to feeding, locomotion, and life history strategy were taken. Findings revealed that taxonomic and functional diversity values changed with invasion in both tests. Taxonomic diversity was 15% more in invaded than uninvaded sections in Test 1 and was not consistent across sampling points of invaded section in Test 2. Invaded areas were taxonomically less diverse, but functionally diverse in both tests. The analysis of similarity and nonmetric multidimensional scaling revealed no difference in Bray–Curtis similarity assemblages in both tests. Our findings revealed that exotic species more often occupy unfilled gaps in the communities often occupied by the native species; this is achieved by occupying functional spaces. Overall, changes in taxonomic and functional diversity of native species documented here partially confirmed impacts of O. niloticus invasion. Therefore, we recommend a multifaceted approach to assess cumulative impacts of invasion on native species.     

Determining the foraging range and origin of resurgence after treatment of Argentine ant (Hymenoptera: Formicidae) in urban areas

The foraging range and distribution of Argentine ant, Linepithema humile (Mayr), colonies in urban areas of southern California extended at least 61 m (200 feet) from feeding stations and structures. Ants were fed at 25% sucrose feeding stations containing 0.01% fluorescent brighter (FB28). Within 14 d, from 77-90% of the ants sampled next to the feeding stations were positive for FB28. The percentage of ants with FB28 declined gradually to approximately 55% 61 m away from the feeding station. The percentage of marked ants in the controls didn't change over the 4-wk-test period. There were approximately 290,000 ants visiting the monitored stations each night before treatments. The 0.0001% fipronil baits and 0.06% fipronil SC sprays provided significant reductions in at least 4 wk. The percentage of ants marked with FB28 decreased significantly in both bait and spray treatments over 4 wk suggesting that the resurgence of ants in treated areas were because of immigration from untreated areas. It is likely that much larger areas will need to be treated to control Argentine ants in urban settings, especially if baits are being used.