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.     

Life-history traits and parasitism rates of four phorid species (Diptera: Phoridae), parasitoids of Atta vollenweideri (Hymenoptera: Formicidae) in Argentina

Leaf-cutting ants in the genus Atta F. (Formicidae, Attini) are among the most important pest arthropods in Central and South America, consuming more vegetation than any other animal group. Among the organisms attacking ants in nature, flies of the family Phoridae have been proposed as the most promising biocontrol agents for pest ants. Four phorid species, Apocephalus setitarsus Brown, Myrmosicarius brandaoi Disney, Myrmosicarius gonzalezae Disney, and Eibesfeldtphora trilobata Disney, were reared from ants collected at Atta vollenweideri Forel nests and off foraging trails in Santa Fe province in Argentina. E. trilobata attacked larger ants and had bigger adults than the other species, also exhibiting the longest developmental time. Correlations between size of hosts and size of adults, as well as between size of adults and developmental times, could be established only in some cases. No differences were found between the sizes of the hosts from which males and females emerged. The natural percentage of parasitism varied throughout the seasons and seemed to be influenced by the extreme drought affecting the study site. We discuss why all four species would be suitable candidates for integrating an assemblage of biocontrol agents against A. vollenweideri.     

Phorid flies, Pseudacteon spp. (Diptera: Phoridae), affect forager size ratios of red imported fire ants Solenopsis invicta (Hymenoptera: Formicidae) in Texas

Multiple species of Pseudacteon phorid flies (Diptera: Phoridae) are currently being released throughout the southern United States to aid biological control of red imported fire ants, Solenopsis invicta Buren (Hymenoptera: Formicidae). It is anticipated that these flies will interfere with S. invicta foraging, allowing native ant assemblages to outcompete S. invicta for available resources. Numerous studies have shown a decrease in S. invicta foraging intensity when exposed to phorids. This study documents a behavioral change in phorid-exposed S. invicta colonies at a phorid release site in central Texas. Significant differences in forager size ratios were detected between phorid-exposed and phorid-absent colonies. A similar phenomenon was recently documented in the native range of these insects in South America as well. Experimental manipulation of ratios of S. invicta worker sizes has been shown to have important effects on colony success. This newly documented phorid-mediated S. invicta colony-level effect represents a significant shift in S. invicta foraging dynamics and may provide an additional mechanism by which phorids can influence S. invicta populations in their United States range.     

Cytogenetic analysis of three species of Pseudacteon (Diptera, Phoridae) parasitoids of the fire ants using standard and molecular techniques

Pseudacteon flies, parasitoids of worker ants, are being intensively studied as potentially effective agents in the biological control of the invasive pest fire ant genus Solenopsis (Hymenoptera: Formicidae). This is the first attempt to describe the karyotype of P. curvatus Borgmeier, P. nocens Borgmeier and P. tricuspis Borgmeier. The three species possess 2n = 6; chromosomes I and II were metacentric in the three species, but chromosome pair III was subtelocentric in P. curvatus and P. tricuspis, and telocentric in P. nocens. All three species possess a C positive band in chromosome II, lack C positive heterochromatin on chromosome I, and are mostly differentiated with respect to chromosome III. P. curvatus and P. tricuspis possess a C positive band, but at different locations, whereas this band is absent in P. nocens. Heterochromatic bands are neither AT nor GC rich as revealed by fluorescent banding. In situ hybridization with an 18S rDNA probe revealed a signal on chromosome II in a similar location to the C positive band in the three species. The apparent lack of morphologically distinct sex chromosomes is consistent with proposals of environmental sex determination in the genus. Small differences detected in chromosome length and morphology suggests that chromosomes have been highly conserved during the evolutionary radiation of Pseudacteon. Possible mechanisms of karyotype evolution in the three species are suggested.     

Sticky traps for monitoring Pseudacteon parasitoids of Solenopsis fire ants

A newly developed method for passively trapping adult Pseudacteon phorid flies, which parasitize workers of Solenopsis saevissima complex fire ants, is described. Adult flies responding to deployed Solenopsis invicta midden were captured when they landed on a Tanglefoot-coated perch, which is part of the trap. This sampling method provided a uniform, repeatable, and verifiable sample that allowed continuous and simultaneous sampling among locations, which can only be accomplished with other techniques by substantially increasing the number of observers. A field test showed the superior operational efficiency and effectiveness of this method relative to other techniques. These traps have been shown effective in various phorid habitats in Texas and Florida. We expect this trap to also be effective in detecting/monitoring phorid flies in other locations.     

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.     

Dynamic expansion in recently introduced populations of fire ant parasitoids (Diptera: Phoridae)

Combating invasive species requires a detailed, mechanistic understanding of the manner and speed with which organisms expand their ranges. Biological control efforts provide an opportunity to study the process of species invasions and range expansions under known initial conditions. This study examines the rate, pattern and mechanisms of spread for two populations of the biological control agent Pseudacteon tricuspis, phorid-fly parasitoids of imported fire ants. We employ a trap-based survey method that detects phorid flies in low-density populations, and provides data on abundance. This technique allows us to differentiate between continuous population spread and effective long-distance dispersal and to examine density gradients of phorid flies across the expanding population front. We find that occupied sites in front of the leading edge of continuous populations were common; forming small populations we refer to as satellite populations. Satellite populations are tens of kilometers from the nearest possible source. Wind governs the dynamics of spread in these two central Texas populations. Population edges expanding with the wind exhibited a higher frequency of effective long-distance dispersal than did populations expanding into the wind. This enhanced effective long-distance dispersal rate translated into a five times faster rate of spread for population edges traveling with the wind. This planned invasion shares many characteristics in common with unplanned species invasions including: protracted establishment phase during which densities were below detection thresholds, and slow initial spread immediately after establishment followed by rapid, accelerating spread rates as population sizes grew.     

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.     

Characterizing the interaction between fire ants (Hymenoptera: Formicidae) and developing soybean plants

This research characterizes the interaction between the fire ants Solenopsis invicta Buren and developing soybean plants. Phagostimulant studies showed that fire ant foraging on soybean seeds increased once the seeds imbibed water. During seedling development over a 5-d germination period, fire ant foraging shifted from the stem/cotyledons to the roots, despite continual increases in fresh weights for each region, and the fact that stem/cotyledon tissue contained the majority of food reserves. Carbohydrate analysis showed that although 2-d-old seedlings had higher concentrations of phagostimulant carbohydrates, especially sucrose, than tissues of mature plants, all tissues analyzed had enough of these sugars to induce a phagostimulant response. Fire ant association with seeds/seedlings germinated in soil resulted in reduced seedling vigor, as determined by a doubling of seedling emergence time, a threefold increase in malformed seedlings, and visible damage to cotyledons. Seeds germinated and grown to mature plants in association with fire ants, allocated 43% more assimilate into pods, but produced 28% less root dry matter, 11% less total dry matter, and there was an 81% reduction in the number of root nodules compared with control plants. We propose that reduced root development and inhibitions of nodule formation would be major yield limiting factors under field conditions. This work demonstrates that fire ant damage to soybeans is not limited to seedling establishment and that more research should be directed at the subterranean activities of the fire ant.     

Repellency of callicarpenal and intermedeol against workers of imported fire ants (Hymenoptera: Formicidae)

Callicarpenal and intermedeol are two insect-repellent terpenoids isolated from leaves of American beautyberry (Callicarpa americana L.; Verbenaceae) and Japanese beautyberry (Callicarpa japonica Thunb.). The repellency of these two terpenoids against workers of red imported fire ants, Solenopsis invicta Buren, black imported fire ants, Solenopsis richteri Forel, and a hybrid of these two species was evaluated using digging bioassays. In a multiple choice digging bioassay using two colonies from each species and their hybrid, callicarpenal showed significant repellency at concentration as low as 50 ppm against both red imported fire ant colonies and 6.25 ppm against all black imported fire ant and hybrid colonies. Intermedeol showed significant repellency at concentration as low as 1.50 ppm against both red imported fire ant colonies and 6.25 ppm against all black imported fire ant and hybrid colonies. In total, 15 colonies, five colonies from each species and the hybrid, were tested on callicarpenal and intermedeol at 50 ppm in a two-choice digging bioassay. Both callicarpenal and intermedeol showed repellency against all colonies, and intermedeol showed significantly greater repellency than callicarpenal against both species and their hybrid.     

The internal phylogeny of ants (Hymenoptera: Formicidae)

Abstract. The higher phylogeny of the Formicidae was analysed using 68 characters and 19 taxa: the 14 currently recognized ant subfamilies plus 5 potentially critical infrasubfamilial taxa. The results justified the recognition of 3 additional subfamilies: Aenictogitoninae Ashmead (new status), Apomyrminae Dlussky & Fedoseeva (new status), and Leptanilloidinae Bolton (new subfamily). A second analysis on these better delimited 17 subfamilies resulted in 24 equally most parsimonious trees. All trees showed a basal division of extant Formicidae into two groups, the first containing (Myrmicinae, Pseudomyrmecinae, Nothomyrmeciinae, Myrmeciinae, Formicinae, Dolichoderinae, Aneuretinae) and the second the remaining subfamilies. Clades appearing within these groups included the Cerapachyinae plus 'army ants', the Nothomyrmeciinae plus Myrmeciinae, the 'formicoid' subfamilies (Aneuretinae + Dolichoderinae + Formicinae), and the Old World army ants (Aenictinae + Aenictogitoninae + Doryline), but relationships within the last two groups were not resolved, and the relative positions of the Apomyrminae, Leptanillinae and Ponerinae remained ambiguous. Moreover, a bootstrap analysis produced a consensus tree in which all branches were represented in proportions much lower than 95%. A reconstruction of the ground plan of the Formicidae indicated that the most specialized of all recent ants are the members of the subfamily Dorylinae and the least specialized ones are the monotypic Apomyrminae.     

Fly pupae as attractant carriers for toxic baits for red imported fire ants (Hymenoptera: Formicidae)

Eight laboratory-reared ant species were fed baits of house fly, Musca domestica L., pupae treated with hydramethylnon. Two fire ant species, Solenopsis invicta Buren and Solenopsis geminata (F.), and Pheidole morrissi (Forel) were killed; average percentage of mortality of the five other species was less than 20%. In contrast, all species that were fed the commercial fire ant bait containing hydramethylnon (Amdro) died or were adversely affected. In the field, applications of house fly pupae and eye gnat, Hippelates pusio Loew, pupae dipped in acetone solutions of fenoxycarb significantly reduced population indices of the red imported fire ant, S. invicta, compared with commercial formulations of fenoxycarb (Logic) and hydramethylnon (Amdro). Field observations showed that the pupae of either species can be carried or moved by one or two worker ants. The smooth, hard cuticle of the pupae make them easy to handle and apply with application equipment. The current cost of house fly pupae is considerably more than the cost of a granular carrier, pregel defatted corn grits. However, if mass-production methods reduce this price differential, fly pupae could become an effective and more species-specific fire ant bait carrier.     

Repellency and toxicity of mint oil granules to red imported fire ants (Hymenoptera: Formicidae)

Repellency and toxicity of 2% mint oil granules were evaluated against worker red imported fire ants. Solenopsis invicta Buren, in a series of laboratory and field experiments. In continuous exposure experiments, LT50 values ranged from 1.2 h with 164.8 mg/cm2 of 2% mint oil granules to 15.3 h with 1.65 mg/cm2 of granules. LT50 values declined exponentially with increasing rate of mint oil granules. Limited exposure to 164.8 mg/cm2 mint oil granules resulted in > 50% knock down (KD) after 30 min; however, there was no KD at 15 min. Twenty-four hour mortality increased linearly with increasing exposure time. Mean repellency of worker red imported fire ants ranged from 49.2 +/- 5.4% for 0 mg/cm2 (untreated control) of mint oil granules at 30 min to 100% for 147.8 mg/cm2 of mint oil granules at 3 h. Repellency increased with increasing milligrams per square centimeter of mint oil granules and exposure time. In field tests, 100% of mounds opened and treated with mint oil granules were abandoned 5 d after treatment and ants had relocated or formed satellite mounds by 2 d after treatment. Unopened mounds treated topically with mint oil granules were not abandoned, but ants formed satellite mounds 2 d after treatment. Mint oil granules could provide another tool for red imported fire ant integrated pest management, particularly in situations in which conventional insecticides would be inappropriate.     

Parasitism and biology of Myrmosicarius grandicornis (Diptera, Phoridae) in relationship to its host, the leaf-cutting ant Atta sexdens (Hymenoptera, Formicidae)

Summary. The parasitic phorid Myrmosicarius grandicornis Borgmeier is commonly found around nest entrances of the leaf-cutting ant Atta sexdens (L.) in Brazil, but there is no information about the importance of this fly for A. sexdens. We evaluated the parasitic capability of female M. grandicornis collected in the field and released in laboratory nests of A. sexdens and compared ants' foraging rhythm before and after M. grandicornis were released. We also determined biological characteristics of the parasitoid. Presence of M. grandicornis elicited foragers to abandon their loads and return to the nest, an effect previously described for the phorid Neodohrniphora sp., which is a more abundant A. sexdens parasitoid. Both phorid species occur at the same foraging trails during part of the year and attack ants of different size classes. Therefore, they may have a combined effect on the reduction of A. sexdens foraging. Myrmosicarius grandicornis pupariated inside hosts' head capsules, which are detached from their bodies and deposited in the nest's refuse pile. Adults emerged through the hosts' mouth cavity about a month after oviposition. The characteristics of M. grandicornis biology suggest that this parasitoid completes its development inside A. sexdens nests.     

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.