Development of RAPD markers in two introduced fire ants, Solenopsis invicta and S. richteri, and their application to the study of a hybrid zone

We developed RAPD DNA markers useful in distinguishing the fire ants Solenopsis invicta and S. richteri. An initial survey of 200 primers revealed seven informative markers; family studies allowed us to determine expression patterns and to confirm Mendelian inheritance of these markers. The seven RAPD markers, one of which is inherited as a codominant marker, were employed along with three allozyme markers to describe the structure of a hybrid zone that has formed between the two species in the USA, where they have been introduced. The data suggest minor introgression of alleles from one parental species ( S. richteri ) into the other ( S. invicta ), which most likely reflects the documented recent movement of this hybrid zone. This pattern is interpreted as interspecific introgression rather than shared intraspecific polymorphism on the basis of comparisons with samples from native, non-hybridizing populations in South America that lack such polymorphism. The data further reveal that the structure of the hybrid zone in the USA varies geographically. One parental species ( S. invicta ) and the hybrids exhibit a mosaic distribution in the east; a gradual transition between the parental species occurs in the centre, with a large intervening area of hybrid genotypes only; and there is apparent contact between parental populations with a small or no intervening zone of hybridization in the west. These differing patterns in the structure of the hybrid zone presumably reflect the unique histories of colonization in different parts of the range of introduced fire ants.     

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.     

Experimental analysis of territory size in a population of the fire ant Solenopsis invicta

Alternative models of territoriality are based on contrastingassumptions about the behavioral processes determining territorysize. In a series of controlled field experiments on the fireant Solenopsis invicta, I tested whether territory size is affectedby the availability of food, as predicted by most economic models,and whether territory size is affected by fighting ability,as predicted by models of competition among neighbors. Abundantfood was offered for 30–35 days to selected colonies eitherimmediately next to the nest (experiment 1) or at peripheralsites near the territory boundary (experiment 2). These foodsupplements had no detectable effect on territory size. Furthermore,food placed near the periphery of the territory did not significantlyalter local boundary positions. During both experiments, largecolonies lost more territory than did small colonies, reflectingtemporary declines in worker number due to the seasonal productionof reproductives. Such losses by large colonies during the summermonths create opportunities for newly founded colonies to expandterritories. In a third experiment, colonies from which workerswere removed lost significantly more territory than did unmanipulatedcontrols. These results show that territory areas in S. invictaare strongly affected by the relative fighting ability of neighboringcolonies but provide no evidence that colonies adjust territoryarea in response to short-term changes in the availability offood.     

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.     

Social chromosome variants differentially affect queen determination and the survival of workers in the fire ant Solenopsis invicta

Intraspecific variation in social organization is common, yet the underlying causes are rarely known. An exception is the fire ant Solenopsis invicta in which the existence of two distinct forms of social colony organization is under the control of the two variants of a pair of social chromosomes, SB and Sb. Colonies containing exclusively SB/SB workers accept only one single queen and she must be SB/SB. By contrast, when colonies contain more than 10% of SB/Sb workers, they accept several queens but only SB/Sb queens. The variants of the social chromosome are associated with several additional important phenotypic differences, including the size, fecundity and dispersal strategies of queens, aggressiveness of workers, and sperm count in males. However, little is known about whether social chromosome variants affect fitness in other life stages. Here, we perform experiments to determine whether differential selection occurs during development and in adult workers. We find evidence that the Sb variant of the social chromosome increases the likelihood of female brood to develop into queens and that adult SB/Sb workers, the workers that cull SB/SB queens, are overrepresented in comparison to SB/SB workers. This demonstrates that supergenes such as the social chromosome can have complex effects on phenotypes at various stages of development.     

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.     

Population genetic structure of the red imported fire ant, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>, in Taiwan

We generated and analyzed microsatellite genotypic data and mtDNA sequence data from the fire ant Solenopsis invicta collected from two separate infested areas (Taoyuan and Chiayi) in Taiwan to infer the population and colony structure of these recently established populations. These genetic analyses revealed the following patterns: 1) Relatedness among worker nestmates was significantly greater than zero for both social forms from both populations; 2) No significant isolation by distance was found among nests within each social form from either population; 3) Significant mtDNA but no nuclear differentiation occurs between sympatric social forms in Taoyuan; 4) Molecular signatures of genetic bottlenecks associated with recent introductions are evident in both populations; and 5) The two sampled populations, Taoyuan and Chiayi, are highly genetically differentiated at both the nuclear or mtDNA genomes and most likely derive from two separate introductions into Taiwan. While results from these analyses generally were consistent with predictions based on the known biology of these ants and similar studies of S. invicta in the U.S.A. and South America, some patterns likely reflect the recent introduction and human-mediated inadvertent transport of ants in Taiwan. This is the first study to investigate the population and colony structure of fire ants in Taiwan and results from our study represent an important contribution to the ongoing efforts aimed at eradicating this invasive pest in Taiwan. Received 18 June 2007; revised 11 October 2007; accepted 17 October 2007.     

Soil nitrogen availability under grasses of different palatability in a temperate semi‐arid rangeland of central Argentina

Abstract Grazing by domestic livestock is frequently associated with the replacement of high‐nutrient palatable species with low‐nutrient unpalatable species, which may have a substantial effect on nutrient cycling. The objective of the present study was to compare soil N availability and net N mineralization in soils under Poa ligularis (palatable grass) with those in soils under Stipa tenuissima (unpalatable grass) in a temperate semi‐arid rangeland of central Argentina. Nitrogen availability and net mineralization under laboratory and field conditions were measured. Soil N availability under P. ligularis was higher than or similar to soil N availability under S. tenuissima. In situ net N mineralization in the soil under P. ligularis was lower than or similar to net N mineralization in the soil under S. tenuissima. Potential net N mineralization was greater in the soil under P. ligularis than in the soil under S. tenuissima. Our results suggest that the replacement of palatable grasses by unpalatable grasses in the temperate semi‐arid rangelands of central Argentina may imply a reduction in the rate of nutrient cycling.     

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.