Aggressive interactions between Solenopsis invicta and Linepithema humile (Hymenoptera: Formicidae) under laboratory conditions

The Argentine ant, Linepithema humile (Mayr), and the red imported fire ant, Solenopsis invicta Buren, are natural agonists in their country of origin. Since the first report of L. humile in California in 1907 its range expanded statewide, displacing native ant species wherever it spread. Since the discovery of established populations of S. invicta in southern California in 1998, it has been restricted to discrete areas of southern California. However, as these discrete populations expand, they are encountering large populations of L. humile, which are effective competitors for available resources and are particularly aggressive in their encounters with other ant species such as S. invicta. Most Dolichoderine ants such as L. humile do not prefer to forage on baits made with defatted corn grit and soybean oil typically used in red imported fire ant control programs. Applications of these baits in areas where distributions of these species overlap might selectively affect populations of S. invicta and give L. humile a competitive advantage. Three laboratory experiments were conducted to determine the competitive outcomes between S. invicta pitted against L. humile: 1) agonistic behavior of workers in small arenas, 2) colony interactions with different population ratios, and 3) the effects of pyriproxyfen on the competitiveness of S. invicta against L. humile. Populations of S. invicta consisting of major workers killed more L. humile than did minors or a mixture of majors and minors. When paired against L. humile colonies consisting of 1,100 workers, colonies consisting of 38 S. invicta workers were easily defeated by L. humile. Colonies consisting of 450 S. invicta workers plugged their nest entrances, but they were ultimately defeated by L. humile after 13 d. The S. invicta colonies consisting of 1,100 workers took control of the bridge connecting the colonies, invaded the L. humile colony, killed the Argentine ant queens, and removed their brood. Pyriproxyfen-treated fire ant workers took significantly longer to chase the Argentine ants from a connecting bridge than did untreated fire ants. Thus, fire ant baits may have long-term effects on intercolonial aggression between S. invicta and L. humile, especially when Argentine ant populations are high in the summer.     

Red imported fire ant (Hymenoptera: Formicidae) control with a corn grit bait of fenoxycarb without soybean oil

The standard fenoxycarb fire ant bait formulation (Logic), composed of pregel defatted corn grits and soybean oil toxicant, was modified by eliminating the soybean oil. This formulation without soybean oil contained greater than 2 times more fenoxycarb and was as effective as the standard bait formulation against laboratory colonies of red imported fire ant, Solenopsis invicta Buren. In field tests, the modified and standard baits were equally effective in controlling fire ants after 6, 12, and 18 wk. Individual worker ants obtained from plots treated with fenoxycarb baits without soybean oil had greater than 47 times less fenoxycarb than did workers from the plots treated with the standard fenoxycarb baits containing soybean oil.     

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.     

Response of Argentine ants and red imported fire ants to permethrin-impregnated plastic strips: foraging rates, colonization of potted soil, and differential mortality

This study investigated the effects of the permethrin-impregnated plastic on ant mortality and foraging rates, and tested its potential for preventing ants from colonizing potted soil. Direct exposure to the plastic for as short as 1 min caused significant mortality of both red imported fire ants, Solenopsis invicta Buren, and Argentine ants, Linepithema humile (Mayr); however, red imported fire ants were more susceptible than Argentine ants. Knockdown of virtually all ants initially occurred within 15 min after exposure. However, some moribund ants recovered from the effects within 24 h. For example, after 1 min of direct exposure to the permethrin-impregnated plastic, 70% of Argentine ants and 5% of red imported fire ants recovered from the treatment. In established colonies of Argentine ants, significantly fewer ants foraged for food up posts treated with the plastic compared with untreated posts. In addition, colonies responded to introduction of the treatment by significantly reducing their overall foraging rates, even on untreated posts. When pots filled with moistened soil were introduced into established ant colonies, 82% of Argentine ants and 99% of red imported fire ants moved into the soil. In contrast, when a 1-cm-wide coil of the plastic was placed under the pot, no ants moved into the soil. The potential for use of these materials in nursery production is discussed.     

A comparison of floral resource exploitation by native and invasive Argentine ants

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

Disruption of parasitism of the California red scale (Homoptera: Diaspididae) by three ant species (Hymenoptera: Formicidae)

The relative influence of the southern fire ant, Solenopsis xyloni McCook, Argentine ant, Linepithema humile (Mayr), and native gray ant, Formica aerata (Francoeur), on parasitism of California red scale, Aonidiella aurantii Maskell, was studied in the laboratory for two parasitoids, Comperiella bifasciata Howard and Aphytis melinus DeBach. All three ant species reduced percentage parasitism by C. bifasciata and both percentage parasitism and host mutilation by A. melinus. Southern fire ant was the least disruptive and native gray ant the most disruptive. Southern fire ant removed 12% of scale from the lemons, presumably to feed on them, while the other ant species did not exhibit significant removal of scale compared to the controls. Percentage parasitism of California red scale exhibited by C. bifasciata was more than 2-fold the level exhibited by A. melinus. Percentage mutilation of California red scale, including probing and host feeding, was nearly 5-fold higher for A. melinus than C. bifasciata. Because A. melinus required a longer total host examination + oviposition period in the absence of ants than C. bifasciata and because oviposition occurs as the last act in a sequence of behaviors, disruption by the ants had a more significant negative effect on oviposition by A. melinus.     

Detrimental effects of highly efficient interference competition: invasive Argentine ants outcompete native ants at toxic baits

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

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     

Impacts of Argentine ants on avian nesting success

Biological invasions can have severe and widespread impacts on ecological communities. A few species of ants have become particularly damaging invaders but quantitative data of their impacts on many taxa is still lacking. We provide experimental evidence using artificial nests baited with quail eggs that the invasive Argentine ant (Linepithema humile) can be a significant avian nest predator – Argentine ants recruited to more nests and in higher abundance than the native ant species they displace. However, at a site invaded by Argentine ants, we monitored over 400 nests of a ground-nesting species, the Dark-eyed Junco (Junco hyemalis), and found that less than 2% of nests failed as a result of Argentine ant predation/infestation. A review of the literature also suggests that Argentine ants may not be a serious threat to bird nests relative to other predators or parasites. However, invasive ants with the capability of overwhelming prey though stinging (specifically the red-imported fire ant, Solenopsis invicta), may have a higher impact on avian nesting success. Received 14 January 2005; revised 28 April 2005; accepted 12 May 2005.     

Hunger in red imported fire ants and their behavioral response to two liquid bait products

To help manage red imported fire ant, Solenopsis invicta Buren, invasion, several types of pest management systems have been developed, including baits. To accurately test liquid bait effectiveness in the laboratory, we determined that starvation time of 96 h is required for laboratory fire ants to simulate foraging ants in the field. We measured density and viscosity of two commercial baits and 20% sugar water at 25 degrees C and then compared amount of material consumed per ant at these physical properties. Mean densities of 20% sugar water, Dr. Moss, and Terro were 1.051, 1.287, and 1.354 g/ml, respectively, and viscosity of each bait treatment varied in the same order but more drastically (1.7, 32, and 400 centipoises, respectively). Field and laboratory studies demonstrated that bait acceptability may be affected by toxicant and physical properties. Baits that are more dense have more mass per volume and may cause the ant to cease feeding with a lower crop load than when they feed on sugar water. Ants that feed on formulated baits exhibit feeding behaviors different from those that occur when feeding on sugar water. At first glance, one might conclude that the difference is because of the toxicant, but our findings suggest that physical properties of baits may be a factor in this change in feeding behavior.     

Factors governing rate of invasion: a natural experiment using Argentine ants

Predicting the success of biological invasions is a major goal of invasion biology. Determining the causes of invasions, however, can be difficult, owing to the complexity and spatio-temporal heterogeneity of the invasion process. The purpose of this study was to assess factors influencing rate of invasion for the Argentine ant (Linepithema humile), a widespread invasive species. The rate of invasion for 20 independent Argentine ant populations was measured over 4 years in riparian woodlands in the lower Sacramento River Valley of northern California. A priori predictors of rate of invasion included stream flow (a measure of abiotic suitability), disturbance, and native ant richness. In addition, baits were used to estimate the abundance of Argentine ants and native ants at the 20 sites. A multiple regression model accounted for nearly half of the variation in mean rate of invasion (R ² = 0.46), but stream flow was the only significant factor in this analysis. Argentine ants spread, on average, 16 m year⁻¹ at sites with permanent stream flow and retreated, on average, −6 m year⁻¹ at sites with intermittent stream flow. Rate of invasion was independent of both disturbance and native ant richness. Argentine ants recruited to more baits in higher numbers in invaded areas than did native ants in uninvaded areas. In addition, rate of invasion was positively correlated with the proportion of baits recruited to by native ants in uninvaded areas. Together, these findings suggest that abiotic suitability is of paramount importance in determining rate of invasion for the Argentine ant. Received: 16 September 1997 / Accepted: 8 February 1998     

The Effect of Diet and Opponent Size on Aggressive Interactions Involving Caribbean Crazy Ants (Nylanderia fulva)

Biotic interactions are often important in the establishment and spread of invasive species. In particular, competition between introduced and native species can strongly influence the distribution and spread of exotic species and in some cases competition among introduced species can be important. The Caribbean crazy ant, Nylanderia fulva, was recently introduced to the Gulf Coast of Texas, and appears to be spreading inland. It has been hypothesized that competition with the red imported fire ant, Solenopsis invicta, may be an important factor in the spread of crazy ants. We investigated the potential of interspecific competition among these two introduced ants by measuring interspecific aggression between Caribbean crazy ant workers and workers of Solenopsis invicta. Specifically, we examined the effect of body size and diet on individual-level aggressive interactions among crazy ant workers and fire ants. We found that differences in diet did not alter interactions between crazy ant workers from different nests, but carbohydrate level did play an important role in antagonistic interactions with fire ants: crazy ants on low sugar diets were more aggressive and less likely to be killed in aggressive encounters with fire ants. We found that large fire ants engaged in fewer fights with crazy ants than small fire ants, but fire ant size affected neither fire ant nor crazy ant mortality. Overall, crazy ants experienced higher mortality than fire ants after aggressive encounters. Our findings suggest that fire ant workers might outcompete crazy ant workers on an individual level, providing some biotic resistance to crazy ant range expansion. However, this resistance may be overcome by crazy ants that have a restricted sugar intake, which may occur when crazy ants are excluded from resources by fire ants.     

不同浓度茚虫威对红火蚁的防治效果

为探索适合防治红火蚁的茚虫威浓度,利用丙酮溶解茚虫威原药,使用雾化手段与载体和配方其他试剂充分混合,干燥后保存,进行野外试验测定其防治红火蚁的药效。结果表明,研制的0.025%茚虫威饵剂综合防治效果达85.30%,0.05%茚虫威饵剂达100%,0.1%茚虫威饵剂达100%。3个浓度的茚虫威饵剂拥有良好的防治红火蚁药效,0.05%浓度最适合研制红火蚁饵剂。     

Red imported fire ants (Hymenoptera: Formicidae) cover inaccessible surfaces with particles to facilitate food search and transportation

Eusocial insects have evolved diverse particle-use behaviors. A previous study reported that red imported fire ants, Solenopsis invicta Buren, deposited soil particles on substances treated with essential balm, a fire ant repellent. We hypothesized that S. invicta modifies inaccessible surfaces by covering them with soil particles to facilitate food search and transportation. Here, laboratory experiments were conducted to study the particle-covering behavior of S. invicta in response to viscose surfaces or surfaces treated with essential balm or liquid paraffin in the presence of real food (sausage) or non-food objects (acrylic plates). S. invicta workers deposited significantly more soil particles on these three types of treated surfaces than on untreated surfaces. In addition, significantly more particles were relocated on viscose and paraffin-smeared surfaces in the presence of food than in the presence of non-food objects. The particle-covering behavior on viscose surfaces was also observed in the field. Interestingly, when no soil particles were available, ants searched and transported food on viscose surfaces only if the surfaces were artificially covered with sufficient quantities of soil particles but could not do so on viscose surfaces without soil particles or with insufficient quantities of soil particles. In addition, ants actively relocated particles to cover viscose surfaces if the transportation distance was within 200 mm, whereas significantly fewer particles were relocated at longer transportation distances (400 mm). Our study provides a novel example of particle use by fire ants during foraging.     

Assessing the effects of sodium on fire ant foraging in the field and colony growth in the laboratory

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Floral visitation patterns of two invasive ant species and their effects on other hymenopteran visitors

Abstract.  1. Floral nectar of the native Hawaiian 'ōhi'a tree, Metrosideros polymorpha , is an important food source for several native honeycreepers and yellow-faced bees, Hylaeus spp., but is also attractive to invasive ants.
2. I undertook this study to compare floral visitation patterns of two widespread invasive ants, the Argentine ant, Linepithema humile , and the big-headed ant, Pheidole megacephala , and to determine their effects on nectar volume and floral hymenopteran visitors.
3. In the first year of the study, Argentine ants visited inflorescences more frequently than big-headed ants at mid-day and in the afternoon, but did not occur in higher densities than big-headed ants at any time of day. In the following year, Argentine ants visited inflorescences both more frequently and in higher densities than big-headed ants. Argentine ant density had a stronger association with nectar concentration than big-headed ant density.
4. Nectar volume did not differ between ant-excluded and ant-visited inflorescences for either ant species. However, ant density was negatively associated with nectar volume for both species.
5.  Hylaeus spp. never visited inflorescences with big-headed ants, while non-native honeybees visited inflorescences with and without ants of either species in equal frequency.
6. Most studies of the effects of invasive ants on native arthropods have focused on interactions on the ground. Flowers should not be overlooked as microhabitats from which native arthropods may be displaced by invasive ants.     

Interference competition by Argentine ants displaces native ants: implications for biotic resistance to invasion

The Argentine ant Linepithema humile (Dolichoderinae) is one of the most widespread invasive ant species in the world. Throughout its introduced range, it is associated with the loss or reduced abundance of native ant species. The mechanisms by which these native species are displaced have received limited attention, particularly in Australia. The role of interference competition in the displacement of native ant species by L. humile was examined in coastal vegetation in central Victoria (southeastern Australia). Foragers from laboratory colonies placed in the field consistently and rapidly displaced the tyrant ant Iridomyrmex bicknelli, the big-headed ant Pheidole sp. 2, and the pony ant Rhytidoponera victoriae from baits. Numerical and behavioural dominance enabled Argentine ants to displace these ants in just 20 min; the abundance of native species at baits declined 3.5–24 fold in direct relation to the rapid increase in L. humile. Most precipitous was the decline of I. bicknelli, even though species in this typically dominant genus have been hypothesized to limit invasion of L. humile in Australia. Interspecific aggression contributed strongly to the competitive success of Argentine ants at baits. Fighting occurred in 50–75% of all observed interactions between Argentine and native ants. This study indicates that Argentine ants recruit rapidly, numerically dominate, and aggressively displace from baits a range of Australian native ant species from different subfamilies and functional groups. Such direct displacement is likely to reduce native biodiversity and indirectly alter food web structure and ecosystem processes within invaded areas. Biotic resistance to Argentine ant invasion from native ants in this coastal community in southeastern Australia is not supported in this study.     

Improving liquid bait programs for Argentine ant control: bait station density

Argentine ants, Linepithema humile (Mayr), have a positive effect on populations of mealybugs (Pseudococcus spp.) in California vineyards. Previous studies have shown reductions in both ant activity and mealybug numbers after liquid ant baits were deployed in vineyards at densities of 85-620 bait stations/ha. However, bait station densities may need to be <85 bait stations/ha before bait-based strategies for ant control are economically comparable to spray-based insecticide treatments-a condition that, if met, will encourage the commercial adoption of liquid baits for ant control. This research assessed the effectiveness of baits deployed at lower densities. Two field experiments were conducted in commercial vineyards. In experiment 1, baits were deployed at 54-225 bait stations/ha in 2005 and 2006. In experiment 2, baits were deployed at 34-205 bait stations/ha in 2006 only. In both experiments, ant activity and the density of mealybugs in grape fruit clusters at harvest time declined with increasing bait station density. In 2005 only, European fruit lecanium scale [Parthenolecanium corni (Bouché)] were also present in fruit clusters, and scale densities were negatively related to bait station density. The results indicate that the amount of ant and mealybug control achieved by an incremental increase in the number of bait stations per hectare is constant across a broad range of bait station densities. The results are discussed in the context of commercializing liquid ant baits to provide a more sustainable Argentine ant control strategy.     

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.     

红火蚁入侵强度对本地蚂蚁群落物种的发现-支配权衡的影响

【目的】发现-支配权衡(discovery-dominance trade off)在物种间的竞争起着重要作用。本研究旨在了解不同强度的红火蚁Solenopsis invicta入侵对本地蚂蚁群落物种的发现-支配权衡的影响,从而探讨红火蚁与本地蚂蚁的竞争机制。【方法】于2017年10-11月,在云南省楚雄州牟定县的一滇橄榄种植基地利用诱饵法和陷阱法结合的方式引诱和收集蚂蚁,观察并记录常见蚂蚁种类的发现能力、招募能力、支配能力、最早发现诱饵的时间段及个体数。【结果】红火蚁的入侵显著影响了本地蚂蚁群落物种的发现-支配权衡。强入侵区(5.6个活动蚁巢/100 m²)蚂蚁群落中物种的相对发现能力和相对支配能力不存在相关性;弱入侵区(0.14个活动蚁巢/100 m²)蚂蚁群落中两者存在较弱的负相关性,但不显著;对照区(无蚁巢)中两者存在显著负相关性(强入侵区:r=-0.01,P=0.97;弱入侵区:r=-0.60,P=0.21;对照区:r=-0.81,P=0.04)。强入侵区的红火蚁种群普遍在5 min之内就能发现诱饵,而弱入侵区的种群需要10～60 ...