Multivariate analyses of the factors affecting the distribution,abundance and social form of Louisiana fire ants, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>

We surveyed 165 sites to determine the ecological factors influencing the distribution, abundance, and occurrence of polygyny in the red imported fire ant (Solenopsis invicta) in Louisiana. On average, sites had 220 nests/ha, 14% of mounds were polygyne, and 22% of sites had ≥ one polygyne mound. The density of nests and ants per site both increased with the proportion of mounds that were polygyne and the organic and phosphorous content of the soil but decreased with longitude, latitude, and the silt: clay, calcium and sodium content of the soil. Ant density also declined with ambient relative humidity. These multivariate models explained ~25% of the variation in nest and ant density per site. Mean mound size per site increased with the phosphorous content of the soil and the number of nests at the site suggesting that prospective queens may select sites that are conducive to produce large mounds. Mean nest size, however, decreased with the proportion of nests that were polygyne and soil potassium while mounds in forests were typically larger than those in residential areas. Overall, this model accounted for 29% of the variation in mean nest size per site. Polygyne sites were patchily distributed across Louisiana. The probability of a site being polygyne declined with mean monthly temperature for 1999 – 2003 and distance to the nearest commercial waterway suggesting that shipping activities may have played a role in the introduction of polygyne colonists to an area. Forested sites were also less likely to be polygyne than those in residential areas. Finally, the density of polygyne nests and ants increased with latitude whereas that of the monogyne form generally declined with latitude. The abundance of both social forms was also greater when they occurred alone. These data are consistent with the hypothesis that monogyne and polygyne S. invicta compete with one another. Received 28 July 2006; revised 2 March 2007; accepted 29 May 2007.     

Nest structure and colony cycle of the Allegheny mound ant,Formica exsectoides Forel (Hymenoptera: Formicidae)

Summary The nest structure and colony cycle of a population of Allegheny mound ants,Formica exsectoides, were examined in central Michigan. The dispersion pattern of mounds was random. Nest structure and presence of brood were primarily determined by excavation of twenty-three nests over three intervals from June through September. Additional excavations of five nests in 1990 and ten nests in 1991 provided further details on nest structure and colony cycle. Most galleries occurred within the mound and upper 30 cm of soil, but some activity reached depths of 100 to 270 cm. Depth of nests showed little correlation with external measurements of height and diameter. Immature stages were recovered from two strata: the upper 20 cm of nest and mound and the lowest nest depths. Alate sexual forms were found in or near the mound in July, and numerous dealate queens were collected in September from peripheral galleries near the soil surface.     

Influence of mound construction by red and hybrid imported fire ants on soil chemical properties and turfgrass in a sod production agroecosystem

Mound construction by imported fire ants (Solenopsis spp.) actively modify the biogeochemical and physical properties of soil; however, their influence on nutrient levels in surrounding vegetation is poorly understood. Aside from the reported persistence of elevated available P and K levels in clay-rich soils one year after mound abandonment, the relative stability of nutrient concentrations from one season to the next is largely unknown. Nutrient concentrations were concurrently analysed from ant mounds and undisturbed soils as well as plant samples collected from warm-season turfgrass in a commercial sod production agroecosystem. Initial collection of soil and turfgrass samples coincided with peak annual biomass (September 2006); the second soil sample collection occurred over twelve weeks later during turfgrass dormancy and ant brood minimum (December 2006). Total C, C/N ratios, organic matter (OM), and Zn²⁺ concentrations as well as pH of ant mound soils were significantly higher than control plot soils; these trends persisted across seasons. Turfgrass harvested from ant mound perimeters in September exhibited elevated N, P, Ca²⁺, S, Cu²⁺, Fe²⁺, and Na⁺ concentrations. Evaluation of the relative stability of soil parameters across seasons revealed a significant drop in ant nest pH from September to December 2006. Total N of mound soils was distinctively greater than control soil counterparts during September only. Soil P, K⁺, Ca²⁺, Mg²⁺ and S (all macronutrients), as well as Na⁺ concentrations from ant mound soils were substantially elevated during the late Autumn to Winter transition compared to control soil locations, whereas Fe²⁺ and Mn²⁺ levels (both micronutrients) were significantly lower in ant mound soils versus control soil environments. Continuous pedoturbation by imported fire ants as well as seasonal shifts in mound soil chemistry resulting from changes in assimilation/dissimilation among mound biota may influence the site-specific effectiveness of microfaunal pathogens (e.g., Thelohania solenopsae) or parasites (e.g., Orasema spp.) identified as classical biological control agents of non-native Solenopsis spp. Therefore, further study of the intrinsic complexities of soil ecosystem dynamics of imported fire ant mounds across several seasons is warranted. Received 10 September 2007; revised 23 April 2008; accepted 1 May 2008.     

Nest mounds of red wood ants (Formicaaquilonia): hot spots for litter-dwelling earthworms

A previously undocumented association between earthworms and red wood ants (Formicaaquilonia Yarr.) was found during an investigation of the influence of wood ants on the distribution and abundance of soil animals in boreal forest soil. Ant nest mounds and the surrounding soil of the ant territories were sampled. The ant nest mound surface (the uppermost 5-cm layer) harboured a much more abundant earthworm community than the surrounding soil; the biomass of the earthworms was about 7 times higher in the nests than in the soil. Dendrodrilusrubidus dominated the earthworm community in the nests, while in soils Dendrobaenaoctaedra was more abundant. Favorable temperature, moisture and pH (Ca content), together with abundant food supply (microbes and decomposing litter) are likely to make a nest mound a preferred habitat for earthworms, provided that they are not preyed upon by the ants. We also conducted laboratory experiments to study antipredation mechanisms of earthworms against ants. The experiments showed that earthworms do not escape predation by avoiding contact with ants in their nests. The earthworm mucus repelled the ants, suggesting a chemical defence against predation. Earthworms probably prevent the nest mounds from becoming overgrown by moulds and fungi, indicating possible mutualistic relationships between the earthworms and the ants. Received: 21 November 1996 / Accepted: 3 April 1997     

Eradication of Solenopsis invicta by pyriproxyfen at the Shihmen Reservoir in northern Taiwan

Abstract The aim of this study was to investigate the possibility of eradicating the red imported fire ant, Solenopsis invicta Buren, in the hot, humid and rainy conditions of northern Taiwan. Thirteen hectares of the Shihmen Reservoir have been infected with these ants, with a total of 1 578 mounds. Pyriproxyfen was applied at a rate of 2 kg/hectare each season, for a total of four seasons. Surveys using visual examination, bait traps, as well as the population index method showed that the brood, the reproductive ants, and the worker ants within the nests were all significantly reduced 1 month after the first application of pyriproxyfen. Four months after the initial application there were no more brood or reproductive ants. After 8 months there were no longer any active mounds left, and 12 months later there were no more worker ants. After monitoring for 1 more year, it was ascertained that the red imported fire ants were eradicated. The percentages of decrease after 2, 4 and 6 months of treatment were 57.4%, 80.5%, and 98.2%, respectively.     

Advancement on techniques for the separation and maintenance of the red imported fire ant colonies

Advancement has recently been made on the techniques for separating and maintaining colonies of red imported fire ants, Solenopsis invicta Buren. A new brood rescue method significantly improved the efficiency in separating colony from mound soil. Furthermore, a new method was developed to separate brood from the colony using fire ant repellants. Finally, a cost-effective method was developed to coat containers with diluted Fluon （AGC Chemicals America, Inc, Moorestown, NJ, USA）, an aqueous polytetrafluoroethylene, to prevent housed ants from escaping a container. Usually the original Fluon solution is directly applied to the wall of the containers. Reduced concentrations of Fhion were found to be equally effective in preventing ant escape. The use of diluted Fluon solutions to coat the containers was recommended because of environmental and cost-saving benefits. Application of these new techniques can significantly reduce labor, cost and environmental contamination. This review paper collates all the new techniques in one reference which readers can use as a manual.     

The economics of brood raiding and nest consolidation during ant colony founding

Summary The most dangerous time for an ant colony is during the founding stage when the small colony is vulnerable to predation and competition. Colonies can grow more rapidly when multiple queens cooperate in raising the first worker brood (pleometrosis) or by raiding other incipient colonies for their brood. This brood raiding has been proposed to be the primary force selecting for pleometrosis, i.e. multiple-queen colonies may have a considerable advantage in destroying neighbours by aggressively stealing their brood. An alternative hypothesis is that incipient nests are part of a larger, interconnected population structure and that brood raiding reflects cooperative pleometrosis with subdivided colonies. A simple mathematical model supports the second hypothesis: workers of incipient colonies are especially favoured to peaceably abandon their nest and join with other colonies if the queens are related or queens from raided colonies can infiltrate the raiding colony. The latter condition is often met in ant species that brood raid and particularly exemplified in fire ants (Solenopsis invicta), where brood raiding involves little mortal combat and combines with pleometrosis to rapidly increase colony size. It is proposed that the term nest consolidation should replace brood raiding to more accurately reflect the relatively non-aggressive and potentially apparently cooperative nature of interactions between incipient ant colonies.     

Zombie fire ant workers: behavior controlled by decapitating fly parasitoids

Laboratory observations were conducted on four separate red imported fire ant, Solenopsis invicta, colonies that contained workers parasitized by the decapitating fly, Pseudacteon tricuspis. Parasitized S. invicta workers remained inside the nest during parasitoid larval development and left the nest approximately 8 – 10 hours before decapitation by the parasitoid. When parasitized ants left the nest, they were highly mobile, were responsive to tactile stimuli, and showed minimal defensive behavior. Ants ultimately entered into a grass thatch layer, where they were decapitated and the fly maggots pupariated. This study reveals that parasitized ants exhibit behaviors that are consistent with host manipulation to benefit survival of the parasitoid. Received 9 November 2006; revised 26 January 2007; accepted 7 February.     

The effects of the red imported fire ant on seed fate in the longleaf pine ecosystem

The natural patterns of myrmecochory are disrupted by the dominance of red imported fire ants (Solenopsis invicta) in the southeastern United States. This leads to questions about the role of fire ants as seed dispersers. We examined the fate of ant-dispersed seed in the longleaf pine ecosystem. First, we determined removal rates for a suite of common ground cover species. Then, we verified the final location of removed seeds by using a wax cast to examine nest contents, and locating dyed seeds deposited in trash piles on the ground surface. Finally, we determined if the germination rate of seeds deposited by fire ants was affected by ant dispersion. Fire ants were most attracted to elaiosome-bearing seed and collected nonelaiosome-bearing seed at a much lower rate. No seeds were found in the contents of wax castings of fire ant nest chambers, suggesting that seed is not stored within the nest. Of the dyed seeds that we presented to fire ants, 30–40 % were recollected in surface trash piles in the mound vicinity within 1 week following removal. Undiscovered seeds were considered destroyed or buried in foraging tunnels. A small percentage of the deposited seeds were able to germinate, but there was no difference in the percent germination between seeds manipulated by fire ants and the control. Low germination was likely due to a high percentage of immature seeds used in the study. Our findings support a growing body of evidence that fire ants facilitate the movement of seeds in the longleaf pine ecosystem.     

三种植物源药剂对红火蚁的防治效果评估及其对本地蚂蚁群落影响

红火蚁Solenopsis invicta Buren是一种危险性入侵害虫,可对人类健康、农林业生产、公共安全和生态系统产生严重的负面影响。目前,化学农药是防治红火蚁的主要方法。然而,大多数化学农药并不适用于有机农场、水产养殖区等生态敏感区。为筛选出用于生态敏感区红火蚁防治的药剂,本文结合室内和田间试验评估了植物源杀虫剂苦参碱Matrine、鱼藤酮Rotenone和除虫菊素Pyrethrin对红火蚁的防治效果。室内试验结果表明,苦参碱、鱼藤酮和除虫菊素对红火蚁均具有较高的毒性。田间试验表明,25.00 mg/L和12.50 mg/L苦参碱、250.00 mg/L 和125 mg/L鱼藤酮、75.00 mg/L和37.50 mg/L除虫菊素灌巢处理,红火蚁蚁巢减退率分别达到100%、81.25%、87.50%、79.17%、71.73%和66.67%。然而,除25.00 mg/L苦参碱外,其他处理均导致在被灌巢的蚁巢周围短期内出现新蚁巢。灌巢处理后,处理区的蚂蚁多样性指数和均匀度指数升高,优势度指数下降。综上所述,苦参碱、鱼藤酮和除虫菊素适合灌巢处理防治红火蚁,其中0.5%苦参碱可溶性液剂能有效降低红火蚁种群数量,使得蚁巢减退率达100%,同时对本地蚂蚁群落没有负面影响,因此该商品制剂在红火蚁防治方面具有较好的开发潜力。     

Interactive effects of soil-dwelling ants, ant mounds and simulated grazing on local plant community composition

Interactions between aboveground vertebrate herbivores and subterranean yellow meadow ants (Lasius flavus) can drive plant community patterns in grassland ecosystems. Here, we study the relative importance of the presence of ants (L. flavus) and ant mounds under different simulated grazing regimes for biomass production and species composition in plant communities. We set up a greenhouse experiment using intact soil cores with their associated vegetation.We found that plant biomass production in the short term was affected by an interaction between simulated grazing (clipping) and ant mound presence. Clipping homogenized production on and off mounds, while in unclipped situations production was higher off than on mounds. During the experiment, these differences in unclipped situations disappeared, because production on unclipped mounds increased. Plant species richness was on average higher in clipped treatments and patterns did not change significantly over the experimental period. Plant community composition was mainly affected by clipping, which increased the cover of grazing-tolerant plant species. The actual presence of yellow meadow ants did not affect plant community composition and production.We conclude that the interaction between ant mounds and clipping determined plant community composition and biomass production, while the actual presence of ants themselves was not important. Moreover, clipping can overrule effects of ant mounds on biomass production. Only shortly after the cessation of clipping biomass production was affected by ant mound presence, suggesting that only under low intensity clipping ant mounds may become important determining plant production. Therefore, under low intensity grazing ant mounds may drive the formation of small-scale plant patches.     

Nest architecture in the western harvester ant,Pogonomyrmex occidentalis (Cresson)

Summary The structure and thermal environment produced by the nest cones of the western harvester ant,Pogonomyrmex occidentalis, are investigated. The nest cone is oriented so that the longest slope faces towards the southeast ad the nest entrance faces the southeast. The temperature of the soil was monitored at thirty-six locations within the nest in order to measure the daily temperature change as a function of depth, aspect and radial distance from the center of peak of the cone. The occupancy of the nest cone by workers and by brood was assessed by core samples taken at different times of day in different portions of the cone. The brood are only present during the midmorning sample and are present in greatest numbers on the eastern side of the cone. The adult workers are present in the cone in highest numbers during the midmorning, virtually absent from the cone during mid-afternoon and returning in smaller numbers at dusk, apparently spending the night near the surface on the eastern side of the mound. The eastern and southern portions of the cone are occupied most frequently while the western side of the cone is nearly vacant. The nest cone is an adaptation which magnifies the amplitude of daily temperature fluctation. By moving to appropriate areas of the nest cone, the ants can take advantage of the increased range of temperature.     

Susceptibility and behavioral response of red imported fire ant (Hymenoptera: Formicidae) to selected entomogenous nematodes (Rhabditida: Steinernematidae & Heterorhabditidae).

Pathogenicity of infective juveniles of selected Steinernema spp. and Heterorhabditis spp. toward developing and reproductive stages of the red imported fire ant, Solenopsis invicta Buren, was tested under laboratory conditions. At 10(3)-10(5) infective juveniles per Petri dish, mortality of reproductive larvae, pupae, and alates ranged from 28 to 100% at higher doses after 96 h at 23-25 degrees C. Steinernema carpocapsae All was the most consistent species tested; this nematode caused mortality of fire ant larvae, pupae, and alates of 82-94, 64-96, and 38-99%, respectively. Although not susceptible to nematode infection, worker ants vigorously preened nematodes from brood, alates, and themselves. In a field study, S. carpocapsae (5 x 10(6) and 2 x 10(6) drench, 2 x 10(6) infective juvenile infection) was applied to active fire ant mounds in 3.8-liter suspensions. Hydramethylnon (75 ml), a water drench, a water injection, and untreated fire ant mounds were marked and treated. Overall activity in mounds treated with nematodes of hydramethylnon ranged from 40 to 48%. Satellite mound activity accounted for 32-44% of overall activity in mounds treated with nematodes 2 wk after treatment. However, 6 wk after treatment, activity in mounds treated with hydramethylnon was 44%; activity of mounds treated with nematodes ranged from 52 to 80%. Satellite mound activity accounted for 0-24% of overall activity. Whereas a soil drench of S. carpocapsae showed potential as a control method for the red imported fire ant, colony relocation after nematode treatment could limit overall efficacy unless application techniques are developed to overcome or take advantage of the movement.     

Post-Release Host-Specificity Testing of Pseudacteon tricuspis, a Phorid Parasitoid of Solenopsis Invicta Fire Ants

Inherent in any biological control program is the risk of nontarget effects. Pseudacteon tricuspisBorgmeier, a parasitoid phorid fly, has been introduced to the United States from South America as a potential biocontrol agent of the red imported fire ant, Solenopsis invictaBuren. We conducted tests of host specificity on introduced populations of P. tricuspis, which are attracted to alarm pheromones released by their hosts during events such as mound disturbances and interspecific interactions. We monitored disturbed mounds of S. invicta and its close congener, S. geminata(F.), during the expansion of P. tricuspis across north Florida and after populations had been established for ~3 years. We also tested host acceptance in established populations of P. tricuspis by offering trays containing S. invicta, S. geminata, and 14 additional ant species representing 12 different non-Solenopsis genera. Although P. tricuspiswas commonly observed to hover over and attempt to oviposit on S. invicta, we never observed any parasitization attempts on any other ant species. As predicted by laboratory tests, released populations of P. tricuspis appear to be highly host specific and pose no obvious threat to nontarget species.     

Thermoregulatory trade-offs result from vegetation removal by a harvester ant

Abstract.  1. Analyses of ecological trade-offs help to explain how organisms balance competing demands. Harvester ants ( Pogonomyrmex occidentalis ), are conspicuous residents of shortgrass prairie in western North America; worker P. occidentalis actively clear all vegetation from the immediate vicinity of their large gravel mounds. This study is based on the prediction that vegetation clearing yields a thermal trade-off by increasing soil temperatures; during cool periods the resulting increase in soil temperature opens new time windows for activity, while during hot periods the soil temperature is more likely to exceed the maximum thermal tolerance for this species. To test the hypothesis that daily and seasonal trade-offs in ant activity result from vegetation removal, the effects of experimentally applied shade on activity patterns were measured.
2. Harvester ant activity correlated highly with ground temperature; experimental shading of ant mounds shifted daily activity patterns by lowering ground temperature. Shading in the morning significantly delayed the onset of ant activity by preventing solar warming of the mound. In contrast, mid-day experimental shading prevented elevation of ground temperatures to above 50 °C and allowed ants to remain active when lethally high temperatures would normally force them inside the mound.
3. A model derived from field data predicted surface ground temperature (and therefore ant activity) based on air temperature and solar radiation, under conditions of sun and shade. For each of six seasons modelled, shade removal yielded a net gain of activity time. These results indicate that vegetation removal by harvester ants produces an advantageous thermoregulatory effect by helping to maximise activity time.     

Impact of Formica exsecta Nyl. on seed bank and vegetation patterns in a subalpine grassland ecosystem

The mound building ant Formica exsecta Nyl. is widely distributed in grassland ecosystems of the Central European Alps. We studied the impact of these ants on seed bank and vegetation patterns in a 11 ha subalpine grassland, where we counted over 700 active ant mounds. The mounds showed a distinct spatial distribution with most of them being located in tall‐grass, which was rarely visited by ungulates (red deer; Cervus elaphus L.). Heavily grazed short‐grass, in contrast, seemed to be completely avoided by ants as only few mounds were found in this vegetation type. The species composition of the ant mound and grassland seed banks was quite similar, i.e. from 15 common plant species 12 were found in both seed bank types. We found the same proportions of myrmecochorous seeds in ant mound and grassland soil samples. In contrast, the number of seeds was 15 times higher in mound compared with the grassland soil samples. Also, the vegetation growing on ant mounds significantly differed from the vegetation outside the mounds: graminoids dominated on ant mounds, herbaceous and myrmecochorous species in the grassland vegetation. We found significant continuous changes in vegetation composition on gradients from the ant mound centre to 1 m away from the mound edge. Overall, F. exsecta was found to have a considerable impact on seed bank and vegetation patterns in the grassland ecosystem studied. These insects not only altered grassland characteristics in the close surrounding of their mounds, but also seem to affect the entire ecosystem including, for example, the spatial use of the grassland by red deer.     

The effect of bait air broadcasting by unmanned aerial vehicles on the ant community diversity

To explore the control effect of bait broadcasting by unmanned aerial vehicles (UAVs) on the imported fire ant Solenopsis invicta and evaluate its impact on biodiversity, we tested the drop flow rate and broadcasting evenness of fire ant baits with bait exit hatch sizes, as well as the dynamics of fire ants and other ants when different amounts of bait were used. The amount of bait applied per hectare was 6000–15,000 g in orchards. Fourteen days after broadcasting, the control effect of broadcasting 12,000 g and 15,000 g bait per hectare on active mounds and workers reached 90%, which was significantly higher than that achieved with 6000 g and 9000 g. After the application of bait, the number and species of other ants decreased and then increased; this was reflected in changes in the Shannon–Wiener index (H′), Pielou evenness index (E) and Simpson dominance index (C). The results of this study suggest that broadcasting bait by UAVs can achieve good control of S. invicta, and a reduction in native ant diversity can be achieved in the short term.     

The urban fire ant paradox: native fire ants persist in an urban refuge while invasive fire ants dominate natural habitats

In contrast to the widespread extirpation of native fire ants (Solenopsis geminata) across southern US following the invasion by imported red fire ants (S. invicta), some residential areas of Austin form unexpected refuges for native fire ants. Ironically, these urban environments provide refuges for the native fire ants while adjacent natural habitats have been overrun by invasive fire ants. Resistance to invasive fire ants in these urban areas occurs mainly in older residential properties constructed prior to the S. invicta invasion, while more recent construction has allowed establishment by S. invicta. The invasive ability of S. invicta is often attributed to escape from parasitoids and efficient dispersal of polygyne multiple queen colonies. Here we also show the importance of landscape parameters in the invasion process, where low levels of disturbance and continuous plant cover in older residential areas form possible barriers to colonization. Dense leaf cover (high NDVI) was also found to be associated with native ant refuges. Long term residential land ownership may have resulted in lower recent disturbance levels and increased plant cover that support refuges of native fire ants.     

Organic mound-building ants: their impact on soil properties in temperate and boreal forests

Ants are important components of most soil invertebrate communities, and can affect the flow of energy, nutrients and water through many terrestrial ecosystems. The vast majority of ant species build nests in the mineral soil, but a small group of ants in temperate and boreal forests of Eurasia and North America build large parts of their nests above‐ground using organic materials collected from the surrounding soil. Many studies have shown that ants nesting in mineral soil can affect water infiltration rates, soil organic matter (OM) content, and nutrient cycling, but much less is known on how mound‐building ants influence soil physical and chemical properties. In this paper we summarize what is known on the soil impacts of organic mound‐building ants in temperate and boreal forests, and how these ants could be affected by ecosystem disturbance and future climate change. Much of this information comes from studies on Formica rufa group ants in Europe, which showed that CO₂ emissions and concentrations of C, N, and P are usually higher in ant mounds than in the surrounding forest soil. However, ant mounds are a minor component of total soil C and nutrient pools, but they do increase spatial heterogeneity of soil water and available nutrients. Mound‐building ants can also impact tree growth, which could change the quantity and quality of OM added to soil. Forest management, fire, and projected climate change, especially in boreal forests, could affect mound‐building ant population dynamics, and indirectly, soil properties.     

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.