Queen execution in the Argentine ant, Iridomyrmex humilis

ABSTRACT. Field censuses and laboratory experiments show that in the Argentine ant, Iridomyrmex humilis (Mayr), c. 90% of the queens are executed by workers in May, at the beginning of the reproductive season. The reduction in the number of queens probably decreases the inhibition exerted by queens on the differentiation of sexuals and thus allows the production of new queens and males shortly thereafter. In the laboratory, there was no correlation between the percentage of queens executed and their weight or fecundity. At the time of execution of queens, nearly all queens were of the same age; less than 1 year. Therefore it is not likely that the age of queens plays any role in the choice that workers make in the queens they executed. Execution of these queens results in a heavy energetic cost for the colony which amounts c. 8% of the total biomass. This behaviour of workers executing nestmate queens is discussed with regard to possible evolutionary significance at the queen and worker level.     

Volatile constituents of the Argentine ant, Iridomyrmex humilis

Using combined gas chromatography-mass spectrometry, some 55 constituents of Iridomyrmex humilis have been characterized on the basis of total extraction data, and of data reported for the separate extraction of heads, thoraxes and gasters of this ant. Twenty-seven hydrocarbons—normal, unsaturated, and branched—have been identified. Of these the alkenes (5), (11), (17), and (22) are located primarily, if not wholly, in the heads. Four pyrazines (1), (3), (12), and (21) are present in the heads. The terpenoid, iridomyrmecin (4) is the known major extractive of I. humilis; dolichodial (2) and four related cyclopentanoid monoterpene acids (103), (104), (105), and (106) are now reported from the gasters. The iridoids (2) and (4) are constituents of the anal gland. Three esters—isopropyl myristate, methyl palmitate, and dibutyl phthalate—have been characterized from the total extract. Of these the dibutyl phthalate, mainly found in the thoraxes and gasters, is possibly an artifact. Twelve fatty acids, normal, unsaturated, and branched, and two chlorinated phenols, have also been identified from each extract. The chlorinated phenols are possibly artifacts.     

Trail Pheromone of the Argentine Ant,Linepithema humile (Mayr) (Hymenoptera: Formicidae)

The Argentine ant (Linepithema humile) is recognized as one of the world''s most damaging invasive species. One reason for the ecological dominance of introduced Argentine ant populations is their ability to dominate food and habitat resources through the rapid mobilization and recruitment of thousands of workers. More than 30 years ago, studies showed that (Z)-9-hexadecenal strongly attracted Argentine ant workers in a multi-choice olfactometer, suggesting that (Z)-9-hexadecenal might be the trail pheromone, or a component of a trail pheromone mixture. Since then, numerous studies have considered (Z)-9-hexadecenal as the key component of the Argentine ant trails. Here, we report the first chemical analyses of the trails laid by living Argentine ants and find that (Z)-9-hexadecenal is not present in a detectible quantity. Instead, two iridoids, dolichodial and iridomyrmecin, appear to be the primary chemical constituents of the trails. Laboratory choice tests confirmed that Argentine ants were attracted to artificial trails comprised of these two chemicals significantly more often than control trails. Although (Z)-9-hexadecenal was not detected in natural trails, supplementation of artificial dolichodial+iridomyrmecin trails with an extremely low concentraion of (Z)-9-hexadecenal did increase the efficacy of the trail-following behavior. In stark contrast with previous dogma, our study suggests that dolichodial and iridomyrmecin are major components of the Argentine ant trail pheromone. (Z)-9-hexadecenal may act in an additive manner with these iridoids, but it does not occur in detectable quantities in Argentine ant recruitment trails.     

Gyne development in the Argentine ant Iridomyrmex humilis: role of overwintering and queen control

Abstract A possible stimulatory effect of overwintering on gyne development in Iridomyrmex humilis (Mayr) was investigated. Although gyne-potent larvae are present in the nest throughout the year, small queenless colony fragments composed of freshly overwintered ants (collected in late winter or early spring) produced 3–8 times more gynes than comparable fragments made up of non-freshly overwintered ants (collected at other times of the year). Apparently, this stimulatory effect of overwintering acts on both the developmental potential of larvae and the tendency of workers to rear sexually competent larvae as gynes; queenless colony fragments in which either the brood or workers were freshly overwintered produced more gynes than fragments composed of non-freshly overwintered workers or brood but fewer than fragments containing both brood and workers freshly overwintered. This increased sexualization potential of larvae due to overwintering is enough to overcome weak pheromonal inhibition of queens associated with low queen number; a single queen in a large freshly overwintered colony fragment is insufficient to inhibit gyne development, whereas ten queens are totally inhibitory. In non-freshly overwintered colony fragments one queen is completely inhibitory. Variability in egg developmental potential according to queen age does not appear to play a major role in the seasonal production of gynes, because at least some eggs of very young queens (less than 3 weeks old) are capable of gyne development. In the field this stimulatory effect of overwintering is superimposed on a seasonal fluctuation in the combined strength of pheromonal queen control. In southern France, gynes are produced only in spring where they arise primarily from overwintered larvae just after a sharp drop in queen number, and presumably the total level of inhibitory queen pheromone, due to the massive execution of queens by workers.     

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.     

Horizontal transfer of insecticides in laboratory colonies of the Argentine ant (Hymenoptera: Formicidae)

Five insecticides used by urban pest management professionals for ant control and three experimental insecticides were tested to determine whether these insecticides were horizontally transferred among individuals in colonies of Argentine ants, Linepithema humile (Mayr) (Hymenoptera: Formicidae). Ants were exposed to insecticide-treated sand for 1 min and then placed in a colony of untreated ants. Ants exposed to 20 and 40 ppm fipronil readily transferred the insecticide to other individuals in the colony, resulting in high mortality. Most of the transfer and subsequent mortality occurred within 4 d after exposure to treated ants. The other insecticides were not transferred, and ants exhibited mortality rates similar to that of the controls. Experiments in large foraging arenas demonstrated that necrophoresis was an important behavior facilitating the horizontal transfer of fipronil. When ants contacted contaminated corpses in the process of removing them to refuse piles, they received a lethal dose of fipronil and subsequently died. Fipronil-contaminated dead ants that were placed in the vicinity of the nest resulted in significantly higher mortality than did corpses placed in a distant foraging arena (30 cm away). Most of the dead ants accumulated in the vicinity of the nest rather than in the foraging arena, workers retrieving dead ants to refuse piles from the foraging arena. The position effect of insecticide-contaminated corpses relative to the nest and its implication for Argentine ant control are discussed.     

Increases in densities of the citrus red mite,Panonychus citri [Acarina: Tetranychidae], in association with the Argentine ant,Iridomyrmex humilis [Hymenoptera: Formicidae], in southern California citrus

We monitored the activities of the Argentine ant,Iridomyrmex humilis (Mayr), in 3 citrus orchards during 1984 and 1985. We also monitored densities of citrus red mite,Panonychus citri (McGregor), the adult spider mite destroyer,Stethorus picipes Casey [Col.: Coccinellidae], and the predatory mite,Euseius tularensis Congdon [Acarina: Phytoseiidae] during the same period in ant-present and ant-free plots in each orchard. I. humilis was excluded for at least 9 months from trees pruned 60 cm from the ground following a basal application of 1 or 2 % chlorpyrifos 4 EC. Autumn densities ofP. citri in these ant-free plots were significantly lower than those in the plots infested withI. Humilis. S. picipes appeared to be the most important predator, maintaining lowP. citri densities on citrus in southern California in the absence ofI. humilis activity.S. picipes densities increased numerically in response to the early spring and late autumn increases in the density ofP. citri populations. However, in the presence ofI. humilis activity, effective predation was apparently inhibited. Densities ofE. tularensis were unaffected by the presence ofI. humilis, nor didE. tularensis respond to spring or autumn increases in the density ofP. citri populations in any of the orchards.      

Argentine ant (Hymenoptera: Formicidae) trail pheromone enhances consumption of liquid sucrose solution

We investigated whether the Argentine ant, Linepithema humile (Mayr), trail pheromone, Z9-16:Ald, could enhance recruitment to and consumption of liquid sucrose solutions. All tests were done as paired comparisons with a 10% sucrose solution as food. In the laboratory, mixing 20 microl of a 10-microg/ml solution of the pheromone with 50 microl of the 10% sucrose solution increased the number of ants feeding by >150%. In a field test, we combined the trail pheromone with a 10% sucrose solution in 50-ml vials. These vials were covered with a plastic membrane that has 1.5-mm-diameter holes punched uniformly across its surface. Ants could drink from the holes after the vials were inverted. For half of the vials, 1 microg of the pheromone was put onto the plastic membrane before the vials were filled with a 10% sucrose solution. The remaining vials had no pheromone on the plastic membrane. After 4 h we measured the consumption in each vial. Bait consumption with the pheromone was enhanced by 29%. In a 2nd series of tests, vials were left outside for 24 h. The consumption rate was 33% higher with the pheromone compared with the controls that didn't have pheromone.     

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.     

Lethal and sublethal effects of thiacloprid on non-target carpenter ant,Camponotus japonicus Mayr (Hymenoptera: Formicidae)

To control population of Monochamus beetles that transmit pine wood nematode, Bursaphelenchus xylophilus, a variety of insecticides have been applied to forest ecosystems in Korea. Non-target predatory insects can be directly or indirectly exposed to insecticides. We evaluated potential lethal and sublethal effects of thiacloprid on survival and behavior of carpenter ants, Camponotus japonicus Mayr. Field-collected ants were directly exposed to several food items such as thiacloprid-exposed Monochamus beetles, 10% sugar water with thiacloprid concentrations, and 10% sugar water at group and individual levels. In experiment for groups of individuals, dead beetle bodies generally had possible adverse effects on ants through dietary exposure, because two forager ants were dead or paralysis after they were exposed to thiacloprid-exposed Monochamus beetles. At individual level, dietary exposure to thiacloprid at concentrations of 10 and 50?mg/L was lethal to ants, causing paralysis and impaired walking. Mortality of ant workers was higher in direct or indirect exposure treatments than that in the control. Some ants exposed to thiacloprid showed abnormal behavior within a few days, especially at thiacloprid concentrations of 10 and 50?mg/L. However, some of them sometimes recovered from the abnormal behavior in a day. In consequence, application of thiacloprid in pine forests may disrupt species interaction and foraging behavior of ants, but the effect of thiacloprid through trophallaxis behavior should be further assessed using more extensive colonies composed of queen, workers, brood, and so on.     

Stenamma debile (Hymenoptera, Formicidae): Facultative polygyny, and an extreme 1997 sex ratio

Summary: Monogynous and a high proportion of truly polygynous colonies of Stenamma debile were found in populations of this small and inconspicuous ant in southern Germany. Moreover, dissections revealed that queens of different age may co-occur in one nest. Thus polygyny is not only an outcome of pleometrosis. Probably young queens sometimes join the mother colonies.¶Practically no female sexuals were reared in the 1997 brood of 29 colonies, thus a population-level sex ratio of &gif1; was recorded. It is likely that such an extreme sex ratio is atypical for this species, suggesting substantial year-to-year variation in sex ratios. Variation of sex ratios over several years has been reported for other ant species, too, but rarely to such an extent. The reason for the failure of queen production in 1997 was perhaps an unusual sequence of warm and cold periods in the spring which may have caused a re-determination of the queen-destined hibernated larvae to become workers, while male larvae were reared in ordinary numbers.     

Multiply Mating Males in Gnamptogenys striatula Mayr (Hymenoptera, Formicidae)

Multiply mating ant males are rare, because in most ant species, multiple mating opportunities for males are scarce. Gnamptogenys striatula is an exception to this rule, as intranidal mating offers protection and a local female-biased sex ratio. In a behavioral experiment, we confirm the males’ ability to inseminate several receptive workers, with four females being the observed maximum. D. Allard is a Research Assistant for the Fund of Scientific Research—Flanders.     

Acceptance and intake of gel and liquid sucrose compositions by the Argentine ant (Hymenoptera: Formicidae)

Liquids and gels are common delivery forms used in commercial ant baits, but the relative effectiveness of each is unknown. We compared the feeding responses of the Argentine ant, Linepithema humile (Mayr), to liquid and gel compositions of sucrose. In choice assays, more workers were counted on gel than liquid; however, substantially more liquid was consumed. Because workers could stand on the gel, more workers could feed simultaneously on the gel. The feeding bouts of individual workers, however, were much less efficient at extracting sucrose in gel form. Workers fed eightfold longer on the gel, yet removed fivefold less sucrose than workers feeding on liquid. This potential bias should be considered during attraction and palatability studies that use physically different bait compositions. When the toxicant fipronil was added to the compositions, a greater proportion of the colony died after workers had fed on liquid than gel baits. This finding suggests that liquid formulations may provide more effective control of Argentine ants due to the greater speed and abundance in which it is ingested.     

Effects of aromatic cedar mulch on the Argentine ant and the odorous house ant (Hymenoptera: Formicidae).

In laboratory studies, the Argentine ant, Linepithema humile (Mayr), and the odorous house ant, Tapinoma sessile (Say), avoided aromatic cedar mulch as a nesting substrate. Both ant species were killed when confined with fresh aromatic cedar mulch in sealed containers. However, when confined with cedar mulch that had been aged outdoors for up to 140 d, mortality of L. humile was complete regardless of mulch age, whereas T. sessile mortality declined significantly over the mulch-aging period. Argentine ant susceptibility to aromatic cedar mulch was also greater than that of the odorous house ant when colonies were restricted to mulch in open trays. In addition, commercial aromatic cedar oil was lethal to both ant species. Our results suggest that aromatic cedar mulch may serve as an effective component of a comprehensive urban ant management program.     

Queen-worker conflict over sex ratio: A comparison of primary and secondary sex ratios in the Argentine ant,Iridomyrmex humilis

We compare the primary sex ratio (proportion of haploid eggs laid by queens) and the secondary sex ratio (proportion of male pupae produced) in the Argentine ant Iridomyrmex humilis with the aim of investigating whether workers control the secondary sex ratio by selectively eliminating male brood. The proportion of haploid eggs produced by queens was close to 0.5 in late winter, decreased to less than 0.3 in spring and summer, and increased again to a value close to 0.5 in fall. Laboratory experiments indicate that temperture is a proximate factor influencing the primary sex ratio with a higher proportion of haploid eggs being laid at colder temperatures. Production of queen pupae ceased in mid-June, about three weeks before that of male pupae. After this time only worker pupae were produced. During the period of production of sexuals, the proportion of male pupae ranged from 0.30 to 0.38. Outside this period no males were reared although haploid eggs were produced all the year round by queens. Workers thus exert a control on the secondary sex ratio by eliminating a proportion of the male brood during the period of sexual production and eliminating all the males during the remainder of the cycle. These data are consistent with workers preferring a more female-biased sex ratio than queens. The evolutionary significance of the production of male eggs by queens all the year round is as yet unclear. It may be a mechanism allowing queen replacement in the case of the death of the queens in the colony.     

Long-term field trial to control the invasive Argentine ant (Hymenoptera: Formicidae) with synthetic trail pheromone

Previous short-term experiments showed that trail following behavior of the Argentine ant, Linepithema humile (Mayr) (Hymenoptera: Formicidae), can be disrupted by a high concentration of synthetic trail pheromone component (Z)-9-hexadecenal. In this study, a long-term field trial was conducted in 100-m2 plots of house gardens in an urban area of Japan to see whether the control effect on Argentine ants can be obtained by permeating synthetic trail pheromone from dispensers. The dispensers were placed in the experimental plots during the ant's active season (April-November) for 2 yr with monthly renewal. To estimate Argentine ant population density, foraging activity of Argentine ants in the study plots was monitored by monthly bait surveys. Throughout the study period, Argentine ant foraging activity was suppressed in the presence of the dispensers, presumably via trail forming inhibition. In contrast, the level of foraging activity was not different between treatment and no-treatment plots when the dispensers were temporarily removed, suggesting that treatment with pheromone dispensers did not suppress Argentine ant density in the treatment plots. Population decline may be expected with larger-scale treatment that covers a significant portion of the ant colony or with improvement in the potency of the disruptant.