Indirect effects of phorid fly parasitoids on the mechanisms of interspecific competition among ants

Indirect effects, which occur when the impact of one species upon another requires the existence of an intermediary species, are apparently very common and may be of greater magnitude than direct effects. Behaviorally mediated indirect effects occur when one species affects the behavior of a second, which in turn affects how that species interacts with a third. I studied behaviorally mediated indirect effects on the mechanisms of competition in two congeneric fire ant species in the presence and absence of parasitoid phorid flies, which parasitized only one ant species. In observational and experimental field studies, the presence of native Texas phorid flies in the genus Pseudacteon decreased food retrieval by their host, Solenopsis geminata (F.), by as much as 50%. In the presence of phorid flies, many S. geminata workers assumed a stationary, curled defensive posture and did not forage. Although the phorid parasitoids had a relatively large effect on exploitative competition, there was no measurable effect on interference competition. Fierce interspecific aggression was observed between S. geminata and S. invicta Buren, and the presence of phorids had no effect on the outcome of these contests. The indirect effects of Pseudacteon parasitoids on Solenopsis fire ant resource retrieval appear to be larger than the direct effect of mortality. Some aspects of the foraging behavior of these Solenopsis species may be, in part, evolutionary adaptations to phorid parasitoid pressure. Because of the relatively large indirect effects, South American Pseudacteon phorids may be promising biocontrol agents of imported fire ants, S. invicta, in the USA. In a laboratory study, a single South American Pseudacteon female was able to significantly decrease food retrieval rates of a North American population of the imported fire ant, S. invicta. Received: 11 May 1998 / Accepted: 18 April 1999     

Fire ant alarm pheromone and venom alkaloids act in concert to attract parasitic phorid flies, Pseudacteon spp.

Pseudacteon tricuspis, Pseudacteon obtusus and Pseudacteon curvatus are three species of parasitic phorid flies (Diptera: Phoridae), which have been introduced as classical biological control agents of imported, Solenopsis fire ants (Hymenoptera: Formicidae) in the southern USA. Previous studies demonstrated the behavioral response of P. tricuspis to the venom alkaloids and alarm pheromone of the fire ant, S. invicta. In the present study, we compared the responses of P. tricuspis, P. obtusus and P. curvatus to Solenopsis invicta alarm pheromone, venom alkaloids, or a mixture of both chemicals in four-choice olfactometer bioassays. The main hypothesis tested was that the fire ant alarm pheromone and venom alkaloids act in concert to attract Pseudacteon phorid flies. Both sexes of all three Pseudacteon species were attracted to low doses of the fire ant alarm pheromone or venom alkaloids (i.e. 1 ant worker equivalent) alone. However, the flies were significantly more attracted to a mixture of both chemicals (i.e., 1:1 mixture of alarm pheromone + alkaloids) than to either chemical. The results suggest an additive rather than a synergistic effect of combining both chemicals. Comparing the fly species, P. tricuspis showed relatively greater attraction to cis alkaloids, whereas the alkaloid mixture (cis + trans) was preferred by P. obtusus and P. curvatus. In general, no key sexual differences were recorded, although females of P. tricuspis and P. obtusus showed slightly higher response than conspecific males to lower doses of the alarm pheromone. The ecological significance of these findings is discussed, and a host location model is proposed for parasitic phorid flies involving the use of fire ant alarm pheromone and venom alkaloids as long range and short range attractants, respectively.     

Dispersal and competitive impacts of a third fire ant decapitating fly (Pseudacteon obtusus) established in North Central Florida

Self-sustaining classical biological control agents offer hope for permanent wide-area control of imported Solenopsis fire ants in the United States because escape from abundant natural enemies left behind in Argentina is a likely reason for unusually high fire ant densities in the United States. The fire ant decapitating fly Pseudacteon obtusus Borgmeier (Diptera: Phoridae) was released as a biocontrol agent of the red imported fire ant (Solenopsis invicta Buren) in Gainesville, FL because it is a common parasitoid of this ant in Argentina and because it has a higher propensity of attacking fire ants along foraging trails than the two Pseudacteon species previously released. Field surveys of a rapidly expanding P. obtusus population (8–12 km/yr) proved that this fly was capable of thriving and successfully competing with the much more abundant Pseudacteon curvatus Borgmeier. However, Pseudacteon tricuspis Borgmeier, the first decapitating fly released, was effectively excluded from most sample sites when faced with competition from both P. curvatus and the similar-sized P. obtusus. Despite clear evidence for competitive exclusion, P. tricuspis abundance at sample sites was positively correlated with the abundance of its two competitors—probably because of moderate to strong covariability in the suitability of sample sites for all three congeners. The addition of P. curvatus, the second parasitoid released, increased total parasitism pressure on fire ant populations by about 10-fold. The addition of P. obtusus, the third species, did not measurably improve total guild parasitism rates on imported fire ants in North Central Florida (as assessed by roadside trap counts), but the performance of this species will likely vary with habitat, region, and climate.     

Comparing longevity of Pseudacteon species of different sizes: effect of sugar feeding

Sugar feeding is known to enhance lifespan in many parasitoid species. Several species of phorid flies in the genus Pseudacteon (Diptera: Phoridae) have been introduced in the Southern U.S.A. for the biological control of imported fire ants of Solenopsis species (Hymenoptera: Formicidae). Pseudacteon species are short‐lived flies and little is known about their nutritional ecology. Results from previous studies in our laboratory show that one of the introduced species, Pseudacteon tricuspis Borgmeier, is capable of feeding on sugar sources with a significant increase in lifespan. However, in Pseudacteon phorid flies, the degree to which sugar feeding can enhance the lifespan of other species is not clear, nor is it known whether there is a relationship between body size and longevity. In the present study, the effect of sugar feeding on the survival of three phorid fly species of different body sizes, Pseudacteon cultellatus Borgmeier, Pseudacteon curvatus Borgmeier and Pseudacteon obtusus Borgmeier, is investigated. For all three species, flies continuously provided with 40% or 20% sucrose solution live longer than individuals provided water only or 40% sucrose solution only on the first day of adult life. However, the degree to which sugar feeding enhanced longevity varies by species and is highest for P. obtusus. The data also indicate that longevity in Pseudacteon phorid flies is related to body size: the largest species (P. obtusus) lives significantly longer than the smaller species (P. cultellatus and P. curvatus). These results suggest that adults of all three Pseudacteon species are capable of feeding on sucrose solution and that sugar feeding can enhance their longevity. The significance of these results is discussed in relation to the field performance of Pseudacteon phorid flies as biological control agents of imported fire ants.     

Pseudacteon decapitating flies (Diptera: Phoridae): Are they potential vectors of the fire ant pathogens Kneallhazia (=Thelohania) solenopsae (Microsporidia: Thelohaniidae) and Vairimorpha invictae (Microsporidia: Burenellidae)?

Fire ant decapitating flies in the genus Pseudacteon were tested for their potential as hosts or vectors of two microsporidian pathogens of the red imported fire ant, Solenopsis invicta. Decapitating flies that attacked or were reared from S. invicta workers infected by Kneallhazia (=Thelohania) solenopsae or Vairimorpha invictae were tested for either pathogen by polymerase chain reaction (PCR) tests or visual examination for spores using phase microscopy. Three species of fire ant decapitating flies acquired the pathogen, Kneallhazia solenopsae. K. solenopsae was detected in 58% of pooled samples of Pseudacteon obtusus flies and 44% of pooled samples of Pseudacteon cultellatus that developed in K. solenopsae-infected fire ant workers. K. solenopsae was also found in 17% of pooled samples of field-collected Pseudacteon curvatus. In contrast, the microsporidium V. invictae was not detected in P. obtusus reared from V. invictae-infected S. invicta workers. Neither K. solenopsae nor V. invictae were detected in any of the hovering or ovipositing flies in the laboratory exposures, indicating no mechanical acquisition of the microsporidia occurred during oviposition activity. Greater than 92% of the P. obtusus that developed in K. solenopsae-infected ants survived and emerged as adults, thus indicating no detrimental effects of the microsporidium on pupal development and emergence. These results indicate that Pseudacteon decapitating flies may be able to vector K. solenopsae but not V. invictae among fire ants. Further tests are planned to determine if flies containing K. solenopsae are capable of transferring this pathogen either during oviposition or by being consumed by fire ant larvae.     

Field Release and Establishment of the Decapitating Fly Pseudacteon curvatus on Red Imported Fire Ants in Florida

Decapitating phorid flies in the genus Pseudacteon are being studied as classical biological control agents of the red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae). Pseudacteon curvatus Borgmeier (Diptera: Phoridae) is a small decapitating fly that attacks small fire ant workers. We released a biotype of P. curvatus from Formosa, Argentina, at three sites near Gainesville, FL. Field releases were conducted in the spring and summer of 2003 and monitored monthly. Flies were discovered within 5 weeks at the spring site and then monthly thereafter. By late spring 2004, flies released at this site had expanded 1.6 km both north and south and about 0.8 km westward. Initially, we found no flies from the two summer 2003 releases but we were successful at finding them 8 months after release during spring 2004. This paper documents the first successful release(s) of P. curvatus on red imported fire ants in the United States.     

The influence of ambient conditions and space on the phenological patterns of a Solenopsis phorid guild in an arid environment

We observed the diurnal distribution of a phorid parasitoid guild of Solenopsis fire ants across five sites in an arid region of western Argentina over 17-months. We found a rich assembly of 15 taxa, of which 7 species were found each month of the year and over most times of the day. The majority of species were found most frequently in the evening. A Canonical Correspondence Analysis of the hourly abundances of the flies in relation to field meteorological conditions suggested that two broad groups of species existed, one of which had flight periods associated with hotter, drier conditions than the second. The first group was most commonly represented by Pseudacteon tricuspis, the P. obtusus complex and P. cultellatus, while some members of the second group, such as the P. nocens complex and P. litoralis were the most abundant and commonly found flies. The range of conditions in which these flies were found suggests that all of the common taxa represent populations that might be suitable for introduction into similarly arid environments of Texas.     

Evaluating biological control of fire ants using phorid flies: effects on competitive interactions

The effects of the parasitic phorid fly, Pseudacteon tricuspis Borgmeier, on the competitive interactions between the red imported fire ant, Solenopsis invicta Buren, and a native North American ant, Forelius pruinosus (Roger), were investigated in the laboratory. P. tricuspis is a highly host-specific endoparasitoid of S. invicta workers that is currently being reared and released as a biological control agent of S. invicta in the US. We tested the effect of P. tricuspis on the colony growth rate of S. invicta when S. invicta was forced to compete with F. pruinosus for a protein resource (freeze-killed crickets) in laboratory competition arenas. In addition to colony growth rate, we quantified the effect of the phorid flies on the foraging rate of S. invicta. Though S. invicta significantly reduced its foraging rate in the presence of the phorid flies, we did not detect an effect of the flies on colony growth rate. Possible explanations for these results include behavioral compensation by S. invicta for the presence of the flies. We present these laboratory results in light of a literature search indicating that laboratory tests of biological control agent efficacy are good predictors of field efficacy. We conclude that P. tricuspis alone is unlikely to suppress S. invicta populations in the field by reducing their competitive ability.     

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.     

Ecological dominance of the red imported fire ant, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>, in its native range

Despite the widespread impacts invasive species can have in introduced populations, little is known about competitive mechanisms and dominance hierarchies between invaders and similar taxa in their native range. This study examines interactions between the red imported fire ant, Solenopsis invicta, and other above-ground foraging ants in two habitats in northeastern Argentina. A combination of pitfall traps and baits was used to characterize the ant communities, their dominance relationships, and to evaluate the effect of phorid flies on the interactions. Twenty-eight ant species coexisted with S. invicta in a gallery forest gap, whereas only ten coexisted with S. invicta in a xerophytic forest grassland. S. invicta was the most numerically dominant species in the richest and complex habitat (gallery forest); however it performed better as discoverer and dominator in the simpler habitat. S. invicta was active during day and night. In spite of its poor capacity to discover resources, S. invicta showed the highest ecological dominance and the second-best behavioral dominance after Camponotus blandus. S. invicta won 78% of the interactions with other ants, mostly against its most frequent competitor, Pheidole cf. obscurithorax, dominating baits via mass recruitment and chemical aggression. P. cf. obscurithorax was the best food discoverer. S. invicta won 80% of the scarce interactions with Linepithema humile. Crematogaster quadriformis was one of the fastest foragers and the only ant that won an equal number of contests against S. invicta. The low presence of phorid flies affected the foraging rate of S. invicta, but not the outcome of interspecific interactions. This study revealed that the red imported fire ant ecologically dominated other terrestrial ants in its native range; however, other species were able to be numerically dominant or co-dominant in its presence.     

Establishment and dispersal of the fire ant decapitating fly Pseudacteon tricuspis in North Florida

The decapitating fly Pseudacteon tricuspis Borgmeier was released at eight sites in North Florida between the summer of 1997 and the fall of 1999 as a self-sustaining biocontrol agent of the red imported fire ant, Solenopsis invicta Buren. Several releases used parasitized fire ant workers while most involved adult flies released over disturbed ant mounds. Establishment and dispersal of fly populations were monitored by disturbing about 10 fire ant mounds at each site and then inspecting them closely for hovering flies over a period of about 30 min. Overwintering populations of flies were successfully established at 6 of 8 release sites. Over several years, fly populations at these sites increased to levels as high or higher than those normally seen in their South American homeland. By the fall of 1999, flies had expanded out 1–6 km from five release sites and occupied about 125 km². By the fall of 2000 the five initial release sites plus one new site had fused into one large area about 70 km in diameter. The flies had expanded out an additional 16–29 km and occupied about 3300 km². By the fall of 2001 the flies had expanded out an additional 10–30 km and occupied approximately 8100 km². Fly dispersal was not related to wind patterns in the Gainesville area. Based on the above rates of dispersal and an establishment rate of 66%, we estimate that a state the size of Florida would require 5–10 releases spaced over a 3-year period to cover the state in 6–9 years.     

Parasitoids deter foraging by Argentine ants (Linepithema humile) in their native habitat in Brazil

The Argentine ant, Linepithema humile, has invaded sites across Africa, Australia, Europe, and North America. In its introduced ranges it eliminates native ants and tends agricultural pests. Few studies have examined the ecology of Argentine ants in their native habitat. This study examined the effects of parasitoid flies, genus Pseudacteon, on the foraging behavior of Argentine ants in part of their native range in southern Brazil. Pseudacteon parasitoids commonly attacked Argentine ants, but not other ant species, in daylight at temperatures above 18°C. Argentine ants abandoned food resources and returned underground in the presence of parasitoids. Parasitoid attack rates diminished as Argentine ants retreated underground. Where parasitoids were present, Argentine ants were abundant at food resources only during times of day when parasitoids were inactive. Where parasitoids were absent, Argentine ants were abundant at food resources throughout the day. Overall, the presence of parasitoids explained observed variation in Argentine ant foraging far better than temperature, although temperature had some effect. The results suggest that Pseudacteon parasitoids inhibit the ability of Argentine ants to gather food resources in their native habitat in Brazil. Received: 11 December 1997 / Accepted: 12 June 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.     

Biogeography and macroecology of phorid flies that attack fire ants in south-eastern Brazil and Argentina

Aim Saevissima group fire ants, Solenopsis richteri and S. invicta, have become serious pests when introduced from Argentina and Brazil to other continents. In South America, Solenopsis are distributed across a great variety of habitats and climates. In North America, S. invicta, introduced free of phorids, now ranges from coast to coast in the south. Success in introducing particular Pseudacteon as agents for the biological control of fire ants has varied across climatic zones. We aimed at assembling all the information about fire ant phorids from Argentina and Brazil, to estimate their richness and geographical ranges, to perform a climatic analysis for these distributions, to define groups and climate‐based communities, and to test and elucidate Rapoport's biogeographical rule. Location Argentina and Brazil (South America). Methods From field and museum collections and historical records, we developed a database of fire ant‐specific phorids throughout their known geographical range. A total of 123 sites with values for 15 climatic variables were mapped between 10° and 38° SL and between 35° and 65° C WL for the presence/absence of phorids. We calculated species richness across all sites combined, and for each phytogeographical region, using rarefaction curves, and ICE and Mmean estimators. We calculated mid‐latitudinal points, geographical ranges and areas for each species. The correlation between mid‐latitudinal point and ranges/areas was tested against a null model generated from the randomization of the raw distributional data. We used several types of multivariate analyses to distinguish groups of phorids by phytogeographical regions, hosts and climate, to find gradients of climate throughout the studied area, to define phorid communities in terms of their relationships with gradients of climate, and to test a mechanism for Rapoport's rule. Results Richness estimations using ICE and Mmean estimators were similar or higher than the observed values depending on the phytogeographical region. Cluster multivariate analyses based on climatic, phytogeographic and host data revealed distinct groupings of Pseudacton. The ‘cerrado’ group was confined to tropical savanna areas. A more ‘widespread’ group included ‘Chaco’ and ‘Maritime’ subgroups defined by their respective association with extreme temperatures or precipitation. Ordination multivariate analyses showed (1) two climatic gradients throughout the study area: one of temperature and the other of precipitation, and (2) that climatic variables significantly explained the observed assemblages of phorids. Positive and negative signs of the eigenvalues from the main axes of a canonical correspondence analysis allowed us to define eight communities whose geographical distribution resembled that of phytogeographical regions. We found a significant and positive correlation between geographical areas and mid latitudinal points, and furthermore, the Mantel test based on climatic variables suggested a mechanism for Rapoport's rule applying in the case of Pseudacteon. Main conclusions Pseudacteon species with greater mid‐latitudinal points occupy broader geographical areas and confront more stressful environmental conditions. Because the composition of Pseudacteon communities is largely determined by climatic variables, the correspondence between climates at sites of origin vs. sites of release should be an important consideration in choosing specific phorids for biocontrol efforts.     

Decreased small mammal and on-host tick abundance in association with invasive red imported fire ants (Solenopsis invicta)

Invasive species may impact pathogen transmission by altering the distributions and interactions among native vertebrate reservoir hosts and arthropod vectors. Here, we examined the direct and indirect effects of the red imported fire ant (Solenopsis invicta) on the native tick, small mammal and pathogen community in southeast Texas. Using a replicated large-scale field manipulation study, we show that small mammals were more abundant on treatment plots where S. invicta populations were experimentally reduced. Our analysis of ticks on small mammal hosts demonstrated a threefold increase in the ticks caught per unit effort on treatment relative to control plots, and elevated tick loads (a 27-fold increase) on one common rodent species. We detected only one known human pathogen (Rickettsia parkeri), present in 1.4% of larvae and 6.7% of nymph on-host Amblyomma maculatum samples but with no significant difference between treatment and control plots. Given that host and vector population dynamics are key drivers of pathogen transmission, the reduced small mammal and tick abundance associated with S. invicta may alter pathogen transmission dynamics over broader spatial scales.     

Phenology and parasitism rates in introduced populations of <Emphasis Type="Italic">Pseudacteon tricuspis</Emphasis>, a parasitoid of <Emphasis Type="Italic">Solenopsis invicta</Emphasis>

We documented patterns of seasonal abundance and rates of parasitism in introduced populations of Pseudacteon tricuspis Borgmeier, a phorid parasitoid of the red imported fire ant, Solenopsis invicta Buren. Adult P. tricuspis populations were censused at monthly intervals for 1 year at three sites in northern Florida. Censuses were conducted by aspirating phorids attracted to disturbed S. invicta mounds. Pseudacteon tricuspis adults were present in every month at all sites, although abundances varied greatly among sites and over time. The highest densities of flies (up to 453 censused at 10 disturbed S. invicta mounds in 30 min) were observed in November, and changes in abundance over time were positively correlated among sites. Sex ratios were usually male biased. Parasitism rates were evaluated by collecting workers from field colonies and monitoring them in the laboratory for evidence of parasitism. Parasitism rates were very low – always less than 1%. The average parasitism rate per colony over 16 colonies and 2 years was 0.058%. No pupariation occurred within the first 8 days of collection, suggesting parasitism by P. tricuspis induced behavioral changes in parasitized workers that precluded such workers from our collections. If so, true field parasitism rates may be several times higher than measured here, yet still low in an absolute sense. These low parasitism rates can be reconciled with observed adult phorid densities by considering the large number of host ants present at the study sites.     

Ecological interactions of Pseudacteon parasitoids and Solenopsis ant hosts: environmental correlates of activity and effects on competitive hierarchies

1. Solenopsis (Hymenoptera: Formicidae) fire ants are host to Pseudacteon (Diptera: Phoridae) parasitoids. The activity of S. geminata (F.) hosts and relative abundance of Pseudacteon phorids, along with five environmental variables, were measured at weekly intervals over an 8‐month period at two sites. 2. Pseudacteon relative abundances often varied greatly from week to week, and were only weakly positively correlated with S. geminata activity. 3. A quadratic function of soil temperature was the single best predictor of ant activity at both sites, explaining 32 and 73% of the variation in ant activity. A linear function of soil moisture was the single best predictor of phorid relative abundance at one site (r² = 0.23) whereas no measured variables were significant predictors of phorid relative abundance at the other site. 4. Interspecific interactions at 600 baits were monitored at a third site to document dominance hierarchies and determine whether the presence of Pseudacteon phorids mediated interspecific interactions in their host, S. geminata. 5. Solenopsis geminata was near the top of dominance hierarchies, which did not diverge greatly from a linear pattern. Three species (S. geminata, S. invicta Buren, and Crematogaster laeviuscula Mayr) won the majority of their interspecific interactions and appear to be co‐dominants at this microhabitat‐rich site. 6. Overall, the presence of phorids had no significant effect on the outcome of interspecific contests involving S. geminata and all other ant species grouped together. Phorids may have contributed to some of the S. geminata losses against other co‐dominant species.     

Phorid fly, Pseudacteon tricuspis, response to alkylpyrazine analogs of a fire ant, Solenopsis invicta, alarm pheromone

The phorid fly, Pseudacteon tricuspis Borgmeier, is a parasitoid of the red imported fire ant, Solenopsis invicta Buren. This fly has been reported to use fire ant chemicals, specifically venom alkaloids and possibly alarm pheromone to locate its host. A recent study identified 2-ethyl-3,6-dimethyl pyrazine as a component of the alarm pheromone of S. invicta. To determine the possible involvement of this fire ant alarm pheromone component in mediating fire ant-phorid fly interactions, we tested electroantennogram (EAG) and behavioral responses of P. tricuspis females to the commercially available mixture of 2-ethyl-3,6-dimethyl pyrazine and its 3,5-dimethyl isomer, as well as six structurally related alkylpyrazine analogs at varying doses. Pseudacteon tricuspis females showed significant EAG response to 2-ethyl-3,6(or 5)-dimethyl pyrazine (herein referred to as pheromone-isomer) at all doses, 0.001-10 μg. Among the tested alkylpyrazine analogs, 2,3-diethyl-5-methyl pyrazine showed significant EAG activity at 0.1 and 1 μg. 2,3-dimethyl pyrazine also showed significant EAG activity at 0.1 μg. Results of four-choice olfactometer bioassays demonstrated significant attraction of P. tricuspis females to the pheromone-isomer (2-ethyl-3,6(or 5)-dimethyl pyrazine) at all tested doses (0.01, 0.1, 1 and 10 μg). The analogs, 2,3-diethyl-5-methyl pyrazine and 2,3-dimethyl pyrazine were significantly better than the control at the higher doses (0.1, 1 and 10 μg). The pheromone-isomer was significantly better than both analogs at two doses, 0.1 and 1 μg. These results confirm that the reported fire ant alarm pheromone component plays a role in mediating attraction of phorid flies to host workers. Venom alkaloids were previously shown to attract P. tricuspis; therefore, we propose that fire ant alarm pheromones may act in tandem or synergistically with venom alkaloids to attract phorid fly parasitoids to fire ant workers.     

Tetradonema solenopsis n. sp. (Nematoda: Tetradonematidae) Parasitic on the Red Imported Fire Ant Solenopsis invicta Buren from Brazil

Explorations in Brazil to find parasites of the red imported fire ant, Solenopsis invicta Buren, have uncovered a new species of the Tetradonematidae, Tetradonema solenopsis n. sp. The nematode parasite was fatal to about 25% of the colony. The female nematodes are large and sausage shaped and the males are small and difficult to find, which is typical of the genus. The ant is a new host record for tetradonematids.     

The Role of Habitat in the Persistence of Fire Ant Populations

The association of the exotic fire ant, Solenopsis invicta with man-modified habitats has been amply demonstrated, but the fate of such populations if ecological succession proceeds has rarely been investigated. Resurvey of a fire ant population in a longleaf pine plantation after 25 years showed that the recovery of the site from habitat disturbance was associated with a large fire ant population decline. Most of the persisting colonies were associated with the disturbance caused by vehicle tracks. In a second study, mature monogyne fire ant colonies that had been planted in experimental plots in native groundcover of the north Florida longleaf pine forest had mostly vanished six years later. These observations and experiments show that S. invicta colonies rarely persist in the native habitat of these pine forests, probably because they are not replaced when they die. A single site harbored a modest population of polygyne fire ants whose persistence was probably facilitated by reproduction through colony fission.