Population genetics and history of the introduced fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae), in Australia

Abstract  The red imported fire ant, Solenopsis invicta , a damaging invasive pest, was discovered in February 2001 in Brisbane, Australia at two sites, Fisherman Islands and suburban Richlands-Wacol. Using four microsatellite loci and the protein marker Gp-9 , we compared the two infestations with each other, and with potential source populations in North and South America to better understand the history of their introduction to Brisbane. Based on an analysis of molecular variance, as well as a maximum likelihood tree of colonies from the two Australian sites, we found that the two sites were genetically distinct and were almost certainly introduced separately. All of the colonies at Fisherman Islands were monogynous, headed by a single queen, while the Richlands-Wacol site had a mixture of single-queen monogynous and multiple-queen polygynous colonies. However, the monogynous and polygynous colonies at the Richlands-Wacol site were not genetically distinct from each other, and probably constitute a single, mixed introduction. Based on allele frequencies at the microsatellite loci, and Gp-9 , both Australian infestations were more similar to North American populations than to South American, though the Fisherman Islands infestation was intermediate, making it difficult to assign. Thus, there has been one introduction from either a North or South American monogynous population at Fisherman Islands, and one introduction from a mixed monogynous/polygynous North American population at Richlands-Wacol. These findings have implications for the control of the current infestations, as well as for the quarantine regulations necessary to prevent additional introductions to Australia.     

Development of Pseudacteon nocens (Diptera: Phoridae) on Solenopsis invicta and Solenopsis richteri fire ants (Hymenoptera: Formicidae)

We studied the developmental performance of the large morph of Pseudacteon nocens Borgmeier (Diptera: Phoridae), a prospective biological control agent of imported fire ants (Hymenoptera: Formicidae). We measured selected life history traits of this parasitoid as a function of 1) host species (Solenopsis invicta Buren versus Solenopsis richteri Forel), 2) temperature (22 versus 28 degrees C), 3) source population of the fly (Corrientes and Santiago del Estero, Argentina), and 4) varied size distributions of offered host ants. Developmental periods were influenced by host species, although the populations responded in opposing manners. Developmental times, however, were most strongly influenced by temperature with total developmental periods lengthened by 17-32% at 22 degrees C. Pupal mortality was also significantly lower at this temperature. Although numbers of progeny per female were significantly higher for the Corrientes population, we found no significant differences in progeny per female according to host species. Interestingly, we found that females were larger than males, and flies from Corrientes were larger than those from Santiago del Estero, even after statistical adjustments for host size. The modal frequency of host size elected in all treatment combinations tested was identical (0.6 mm), a size that represented the apparent threshold for producing female progeny. These laboratory and additional field observations demonstrate considerable interpopulational variation in P. nocens and lend further support to the applied approach focusing at the population, as opposed to the species level, with respect to both source and target areas for classical biological control introductions of Pseudacteon flies.     

Unexpected,well-developed nestmate recognition in laboratory colonies of polygyne imported fire ants (Hymenoptera: Formicidae)

Aggression bioassays were used to investigate nestmate recognition in polygyne laboratory colonies of the imported fire ant, Solenopsis invictaBuren. Unlike workers from polygyne field colonies, laboratory-maintained (>10 weeks) workers exhibited well-developed nestmate recognition. As in monogyne colonies of this species, both heritable and environmentally acquired (diet) odors provided recognition cues and were roughly additive in their effect. Within diet treatments, polygyne colonies responded in a graded fashion to polygyne conspecifics, monogyne conspecifics, and heterospecifics (S. richteri Forel),thus suggesting incipient genetic divergence between the two S. invictasocial forms. Hypotheses to account for the acute intraspecific discrimination observed in the laboratory are presented. Empirical testing of these hypotheses will illuminate ecological constraints and proximate mechanisms underlying the reduced intercolony discrimination associated with natural polygyne colonies of this and other ant species.     

Carbon dioxide concentrations in the nests of the mud-dwelling mangrove ant Polyrhachis sokolova Forel (Hymenoptera: Formicidae)

Abstract The nests of the mangrove ant Polyrhachis sokolova are found in soil in intertidal mangrove communities, and are thus inundated at high tides for several hours. Some of the nest galleries are flooded, but others retain air pockets, to which the ants retreat. During and following inundation, we measured carbon dioxide concentrations in air samples collected from different levels in the nests and from artificial 'control' holes in the mud. To account for the relative contribution of different sources of carbon dioxide, we also measured the carbon dioxide production by individual ants (including larvae and pupae) and small samples of mud collected near the ant nests. Nest carbon dioxide concentrations were high (2.5−11%) during and immediately following inundation, but the concentrations in the upper regions of the nest fell as soil water levels receded. However, at depths>10 cm below the level soil surface, the carbon dioxide concentrations remained relatively high and stable (at approximately 2%) over the 11 days between one high tide and the next. The contribution of the mud (and associated microorganisms) to the carbon dioxide concentration in the nests was substantial, and the contribution of the respiration of the ants was approximately 10−15% of the total. The carbon dioxide concentrations in the nests of this species during high tides are among the highest recorded for insect nests, suggesting that these ants may have unusual physiological attributes to match the behavioural and ecological challenges associated with living in the intertidal zone.     

Bionomics of Orasema simplex (Hymenoptera: Eucharitidae), a parasitoid of Solenopsis fire ants (Hymenoptera: Formicidae) in Argentina

Biological characteristics of the parasitoid Orasema simplex Heraty (Hymenoptera: Eucharitidae), a potential candidate for the biological control of fire ants in the United States were investigated. Female survivorship, fertility and oviposition preferences were studied in the laboratory. Naturally parasitized colonies were examined to determine offspring sex ratio, development success and time, and to artificially parasitize healthy ant colonies. In addition, field studies were carried out to establish natural oviposition substrates and adult activity patterns. Orasema simplex female survivorship was 3.6 ± 1.5 days. Newly emerged females contained 613.5 ± 114.0 mature eggs. The adult development success in natural parasitized colonies was 22.2% with a female-biased sex ratio (4:1). The time required from planidia to adult was 29.5 ± 5.4 days. In the field, adults were mostly found around the ant nests at midday. A broad range of plant species was observed as oviposition substrates. The transfer of planidia to healthy ant colonies was achieved but the development success was very low. Orasema simplex appears to have a limited potential as a fire ant biocontrol agent because of cosmetic damage to a wide variety of plants used for oviposition. However, further studies are necessary to evaluate the real damage exerted by oviposition punctures.     

Male Behavior and Regulation of Worker Mating in a Ponerine Ant, Pachycondyla (Bothroponera) sp. (Hymenoptera: Formicidae)

Male behavior and regulation of worker mating were observed in Indonesian Pachycondyla sp. The colonies had some mated workers but only one gamergate (mated and egg-laying workers). Males frequently tried to mate with workers in their natal nests irrespective of worker age and dominance rank, however, workers never accepted mounting of nestmate males. Worker mating occurred only under the gamergateless condition with alien males.     

Swarming, Delayed Sexual Maturation of Males, and Mating Behavior of Fopius arisanus (Sonan) (Hymenoptera: Braconidae)

Fopius arisanus is unusual among hymenopterous parasitoids in males having an obligatory premating period. We confirmed Hagen's (1953) view that sperm takes several days to migrate from the testes to the seminal vesicles. Males mated for the first time only 4 days after emergence, the time that sperm was first ever recorded in the seminal vesicles. In the field, we studied the sexual maturation of F. arisanus males in relation to their behavior. In general, sexually immature males were found in male-only swarms that persisted over several days in host tree canopies. Mature males were usually found in loose aggregations in the vegetation beneath host trees. Females entered these loose aggregations and were mated, whereas they were generally absent from swarms in the canopy. Swarming is therefore apparently not a primary component of the sexual communication system of the species. We describe, for the first time in F. arisanus, the behavioral sequence that accompanies the intersexual communication that leads to mating. Males probably release a volatile chemical that attracts females from a distance, but we have only circumstantial evidence for this. We also postulate that males may form aggregations to amplify the effects of this distance attractant. In the vicinity of males, females release a pheromone that attracts males, a process we demonstrated with female-baited sticky traps.     

Characterization of 24 microsatellite markers in 11 species of fire ants in the genus Solenopsis (Hymenoptera: Formicidae)

The social parasite ant Solenopsis daguerrei infests colonies of several mound-building fire ant species. Twenty-four microsatellite markers were isolated from a repeat-enriched genomic library of S. daguerrei. Eleven loci were polymorphic in this ant with two to six alleles per locus. Expected heterozygosity ranged from 0.0222 to 0.7940 among loci. Most microsatellites amplified successfully across the 11 Solenopsis species tested and will be useful for evolutionary genetic studies in this diverse ant group.     

External and internal structure of Atta bisphaerica Forel (Hymenoptera: Formicidae) nests

Abstract:  This work was carried out to determine the internal and external structures of Atta bisphaerica (Forel) nests. Six nests were excavated and during excavation, all data referring to chambers and tunnels were recorded. Three nests had been internally cement-moulded, which enabled a better view of the chamber and tunnel structures. Atta bisphaerica nests presented a similar structural pattern, varying only in the number of chambers as a function of external mound area. Chambers were spherical with two communication tunnels. Internal tunnels had an elliptical section, sometimes circular, indirectly linked to chambers through ramifications and directly through short tunnels. Entrance holes were linked to the area of highest chamber concentration by tunnels from the elliptical section, which led to the nest in a radial manner. Knowledge of the colony's three-dimensional architecture permits successful application of chemical control processes, reduces the quantity of product applied, and consequently diminishes costs and environmental damage.     

Mating Behavior and Colony Founding of the Slave-Making Ant Formica sanguinea (Hymenoptera: Formicidae)

The mating and postmating behavior of reproductives belonging to two sympatric dulotic colonies of the facultative slave-making ant Formica sanguinea was analyzed in the field. Our observations showed that the European blood-red ant adopts a reproductive behavior similar to the male aggregation syndrome. Newly mated females return to a dulotic colony and often wait for a raid. Following a slave raid is an advantageous strategy to locate and invade host nests and to establish a new dulotic colony. In the laboratory, the following modes of colony founding were studied: independent, adoption, alliance, usurpation, and brood raiding. Independent foundation was possible only when several females were kept together. Alliance was obtained with females of two potential slave species (F. cunicularia, F. rufibarbis). Usurpation and adoption were more frequent in the incipient than in the mature host colonies. Mixed colonies were always obtained after the sack of the host pupae. It seems likely that, rather than conspecific adoption followed by budding, F. sanguinea relies on temporary parasitism to start new colonies.     

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.     

Further records of ants (Hymenoptera: Formicidae) from Iran

Wide ranging surveys of the ant fauna of Iran have enabled us to add a further 30 named species to the country list. A review of almost all the published literature and of photographs of unidentified specimens within the public domain gives a grand total of 248 species, from seven subfamilies and 37 genera. In the majority of instances, our own specimens were compared with type images available from antweb.com. This has led us to propose new or revised status for Cataglyphis turcomanica Crawley 1920, Lepisiota integrisquama (Kuznetsov-Ugamsky, 1929), Lepisiota surchanica (Kuznetsov-Ugamsky, 1929) and Messor obscurior Crawley 1920. We note that the total includes a number of what may be misidentifications and a small number of named species that seem unlikely to occur in Iran.     

Effects of contaminants on bait acceptance by Solenopsis invicta (Hymenoptera: Formicidae)

Three commonly used fire ant baits, Amdro (0.73% hydramethylnon [AI]), Ascend (0.011% abamectins [AI]), and Maxforce (1.0% hydramethylnon [AI]), were exposed to potential, volatile contaminants. The contaminants included the insecticides Orthene Fire Ant Killer (75.0% acephate [AI] ), Cyren (44.6% chlorpyrifos [AI]), and Tempo 2 (24.3% cyfluthrin [AI]); cigarette smoke; gasoline (unleaded, 89 octane); and fertilizer (10-10-10). Fire ant baits previously exposed for 48 h to these contaminants were analyzed using gas chromatography analysis. Orthene Fire Ant Killer, Cyren, Tempo 2, cigarette smoke, and gasoline had volatile components transferred to the baits. Baits exposed to these products were used in a field evaluation of bait acceptance by Solenopisis invicta Buren, the red imported fire ant. Uncontaminated Amdro was significantly preferred by S. invicta over Amdro contaminated by smoke, Cyren, Tempo 2, and gasoline. Uncontaminated Maxforce was significantly preferred over Maxforce contaminated by Tempo 2, Cyren, and gasoline, and uncontaminated Ascend was preferred over Tempo 2- and Cyren-contaminated Ascend. Orthene-exposed Amdro, Maxforce, and Ascend baits, and smoke-exposed Maxforce and Ascend baits were not significantly different from the control. These results indicate that volatile insecticides and products can contaminate fire ant baits. Some insecticides and products, such as gasoline, can significantly affect bait palatability and may adversely impact control.     

Common names for Australian ants (Hymenoptera: Formicidae)

Abstract Most insects do not have common names, and this is a significant barrier to public interest in them, and to their study by non-specialists. This holds for even highly familiar insect groups such as ants. Here, I propose common names for all major native Australian ant genera and species-groups, as well as for many of the most abundant and distinctive species. Sixty-two genera, 142 species-groups and 50 species are given names. The naming system closely follows taxonomic structure; typically a genus is given a general common name, under which species-group and species names are nested.     

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.     

Morphology and number of Ommatidia in the compound eyes of Solenopsis invicta, Solenopsis richteri, and their hybrid (Hymenoptera: Formicidae)

The morphology and number of ommatidia of the compound eyes of Solenopsis invicta, Solenopsis richteri and their hybrid are investigated. Comparisons are made of the morphology and number of ommatidia in S. invicta, S. richteri and their hybrid. Major workers have significantly more facets than minor workers, male reproductive more than female reproductive, and the reproductives more than workers. Workers and reproductives have significantly more ommatidia in S. invicta than in S. richteri and significantly fewer in the hybrid than in either species. The major workers of each species have significantly wider facets than the minor workers, and workers have significantly wider facets than reproductives, which showed no difference in their facet diameter. There are no significant differences when the facet diameter is compared between the members of the castes of different species. These morphological and numerical dissimilarities are discussed in connection with known behavioral differences in the castes.     

Mating Behavior in Halictine Bees (Hymenoptera: Halictidae)

Studied two types of mate-orientation behavior in ♂♂ of the sweat bee, Lasioglossum rohweri: microterritoriality and patrolling. Observations were made in both the laboratory and the field. These ♂♂ behave uniquely for Hymenoptera in that 2–5 of them establish microterritories immediately around the same nest entrance. Some ♂♂ are exclusively patrollers, flying about flowers and nest sites. Rendezvous places (locations where animals are likely to find mates) in bees are: flowers, nesting sites, vegetative parts of plants, nest entrances, air, and nests. Territoriality in male bees may have arisen independently in 7 of the 9 families of bees.