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.     

Ontogeny of the Defensive Stinging Behavior of the Fire Ant, Solenopsis invicta

The fire ant, Solenopsis invicta, appears to deviate from the usual age-related pattern of defensive behavior seen in other social insects; instead of older workers being more defensive than younger ones, they are less so. Here I test this pattern by quantifying changes in the defensive stinging behavior of S. invicta workers of known age. I found defensiveness, measured as both the number of stings delivered and the amount of venom delivered per sting (venom dose), to increase with age initially but then decline after a mid-age peak. This hump-shaped ontogeny may be the result of S. invicta's foraging strategy, which effectively shifts the responsibility of nest defense to workers younger than foraging age. It is S. invicta's mid-aged workers that are the most defensive, probably because they are both physiologically and spatially the most suitable nest defenders.     

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.     

Interactions between ants attendingAphis fabae ssp.cirsiiacanthoidis on thistles and foraging parasitoid wasps

We describe the behavioral interactions between honeydew-collecting workers of the ants Lasius nigerand Myrmica ruginodisand females of three species of aphidiid wasps (Lysiphlebus cardui, Lysiphlebus testaceipes, Trioxys angelicae)foraging for their aphid host, Aphis fabaessp. cirsiiacanthoidis,on thistles. Using field and laboratory experiments, we show that the ant-parasitoid interactions are species specific. Workers of both ant species generally attacked and killed females of T. angelicae,but they ignored those of L. cardui.This pattern was not altered when we anesthetized the wasps slightly with carbon dioxide to reduce their mobility. Prior contacts between L. carduiand either conspecific L. nigerfrom a different nest or workers of a different ant species (M. ruginodis)did not influence L. niger'snonaggressive behavior. The number of aphids parasitized by L. testaceipeswas significantly reduced in aphid colonies attended by L. niger,although this parasitoid was rarely attacked by ants. In encounters between these species of ants and wasps, ant aggression is consistent with differences in wasp behavior. We suggest that, in addition, chemical cues located in the cuticula may enable L. carduito avoid detection by honeydew-collecting ants.     

Host preference in Pseudacteon phorid flies: response of P. tricuspis and P. curvatus to black, red and hybrid imported Solenopsis fire ants in multiple choice bioassays

Host preferences in both sexes of Pseudacteon tricuspis Borgmeier (Jaguariuna biotype) and Pseudacteon curvatus Borgmeier (Formosa biotype) and their relative attraction to the imported fire ants (IFA), Solenopsis invicta Buren (red IFA), Solenopsis richteri Forel (black IFA) and S. invicta × S. richteri hybrids (hybrid IFA) were investigated in two separate experiments utilizing multiple choice flight bioassays. The results of both experiments clearly showed that both sexes of the Jaguariuna biotype of P. tricuspis could distinguish among the three IFA species and demonstrated greater preference for hybrid IFA and red IFA. This conclusion is supported by a variety of data collected on the number of fly visits, attack rate, and hovering duration (Experiment 1), and on the number of trapped flies (Experiment 2), which showed that black IFA is the least preferred of the three species. Similar results were recorded for the Formosan biotype of P. curvatus, although the data were not as strongly conclusive. Females of this biotype spent a significantly greater amount of time in hovering mode over red IFA and hybrid IFA compared to black IFA, but the other data were not significant. The red IFA is the natural host of both phorid fly biotypes and our results suggest that both biotypes may have evolved a specialized relationship with red IFA including an ability to discriminate it from related fire ants. These results are discussed in relation to the possible role of fire ant chemicals in mediating host preferences in phorid flies, contributions of male phorid flies to fire ant biocontrol, and the practical implications of the key findings.     

Predator and parasitoid attacking ant-attended aphids: effects of predator presence and attending ant species on emerging parasitoid numbers

Interaction between a predator and a parasitoid attacking ant-attended aphids was examined in a system on photinia plants, consisting of the aphid Aphis spiraecola, the two ants Lasius japonicus and Pristomyrmex pungens, the predatory ladybird beetle Scymnus posticalis, and the parasitoid wasp Lysiphlebus japonicus. The ladybird larvae are densely covered with waxy secretion and are never attacked by attending ants. The parasitoid females are often attacked by ants, but successfully oviposit by avoiding ants. The two ants differ in aggressiveness towards aphid enemies. Impacts of the predator larvae and attending ant species on the number of parasitoid adults emerging from mummies per aphid colony were assessed by manipulating the presence of the predator in introduced aphid colonies attended by either ant. The experiment showed a significant negative impact of the predator on emerging parasitoid numbers. This is due to consumption of healthy aphids by the predator and its predation on parasitized aphids containing the parasitoid larvae (intraguild predation). Additionally, attending ant species significantly affected emerging parasitoid numbers, with more parasitoids in P. pungens-attended colonies. This results from the lower extent of interference with parasitoid oviposition by the less aggressive P. pungens. Furthermore, the predator reduced emerging parasitoid numbers more when P. pungens attended aphids. This may be ascribed to larger numbers of the predator and the resulting higher levels of predation on unparasitized and parasitized aphids in P. pungens-attended colonies. In conclusion, a negative effect of the predator on the parasitoid occurs in ant-attended aphid colonies, and the intensity of the interaction is affected by ant species.     

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.     

Impacts of Argentine ants on avian nesting success

Biological invasions can have severe and widespread impacts on ecological communities. A few species of ants have become particularly damaging invaders but quantitative data of their impacts on many taxa is still lacking. We provide experimental evidence using artificial nests baited with quail eggs that the invasive Argentine ant (Linepithema humile) can be a significant avian nest predator – Argentine ants recruited to more nests and in higher abundance than the native ant species they displace. However, at a site invaded by Argentine ants, we monitored over 400 nests of a ground-nesting species, the Dark-eyed Junco (Junco hyemalis), and found that less than 2% of nests failed as a result of Argentine ant predation/infestation. A review of the literature also suggests that Argentine ants may not be a serious threat to bird nests relative to other predators or parasites. However, invasive ants with the capability of overwhelming prey though stinging (specifically the red-imported fire ant, Solenopsis invicta), may have a higher impact on avian nesting success. Received 14 January 2005; revised 28 April 2005; accepted 12 May 2005.     

A newly-discovered mode of colony founding among fire ants

Summary Queen ants start new colonies either unassisted by workers (independent founding), assisted by workers from their natal nest (dependent founding), or assisted by the workers of other species (dependent, socially parasitic). The monogyne form of the fire ant,Solenopsis invicta, founds independently in summer, but in the fall it also produces a few sexuals some of which overwinter, then fly and mate in early spring. These overwintered queens lack the nutritional reserves and behaviors for independent colony founding. Rather, they seek out unrelated, mature, orphaned colonies, enter them and exploit the worker force to found their own colony through intraspecific social parasitism. Success in entering orphaned colonies is higher when these lack overwintered female alates of their own. When such alates are present, orphaning causes some to dealate and become uninseminated replacement queens, usually preventing entry of unrelated, inseminated replacement queens. Such colonies produce large, all-male broods. Successful entry of a parasitic queen robs the host colony of this last chance at reproductive success. Only overwintered sexuals take part in this mode of founding.     

Defensiveness of the fire ant, Solenopsis invicta, is increased during colony rafting

Colonies of the fire ant, Solenopsis invicta, can survive flood conditions by forming a raft of ants that floats on the water’s surface until the flood recedes or higher ground is found. Having been forced from the protection of their subterranean nests, rafting colonies are totally exposed and are without retreat. I tested the hypothesis that rafting S. invicta colonies would compensate for their elevated vulnerability by increasing their defensiveness. I measured defensiveness using the amount of venom workers delivered per sting (venom dose), since the repellent effects (i.e., pain and tissue damage) of fire-ant venom are dose-dependent. In the laboratory I assayed colony defensiveness before and after flooding colonies from their nests with water. Colonies were consistently and significantly more defensive while rafting (i.e., each colony’s workers delivered higher venom doses when their colony was rafting than they did when it was assayed pre-flood). The larger venom doses of rafting colonies may reduce their chances of being damaged by encounters with other animals by reducing the duration of such encounters through increased repellency. Encounters with S. invicta during flood conditions have the potential to be unusually dangerous; large concentrations of workers are exposed and available for defense, and they deliver significantly larger venom doses when they sting. Received 29 March 2005; revised 20 June 2005; accepted 24 June 2005.     

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 Bionomics ofCoptera Haywardi(Ogloblin) (Hymenoptera: Diapriidae) and Other Pupal Parasitoids of Tephritid Fruit Flies (Diptera)

The endoparasitoidCoptera haywardi(Ogloblin) (Diapriidae) was discovered in Mexico attacking the pupae of the Mexican fruit fly,Anastrepha ludens(Loew). Typically, parasitoids of Diptera Cychlorrhapha pupae develop as ectoparasitoids and are generalists that attack hosts in a number of families. Aspects of the bionomics ofC. haywardiwere compared to those of two chalcidoid ectoparasitoids,Dirhinus himalayanusWestwood andSpalangia geminaBoucek.C. haywardideveloped in three genera of Tephritidae, but not in species of other families. The two species of chalcidoids developed in all the calypterate and acalypterate hosts to which they were exposed. In an olfactometerC. haywardipreferredAnastrepha suspensa(Loew) pupae, while the chalcidoids preferred the pupae ofMusca domestica L.This preference inS. geminawas diminished in insects that had been reared onA. suspensa. C. haywardioviposited in theA. suspensapupae that had been previously parasitized by the braconidDiachasmimorpha longicaudata(Ashmead). However, it completed development only in unparasitized pupae. Mortality of the primary parasitoid due toD. himalayanuswas approximately two-thirds the mortality inflicted on the host fly.S. geminadid not discriminate between parasitized and unparasitized pupae ofA. suspensaand developed in both.C. haywardiappears to have a more restricted host range relative to chalcidoid pupal parasitoids and this may be due to its endoparasitic development.     

昆明红火蚁种群发生特征

【目的】2013年10月云南省元谋县首次发现红火蚁。掌握红火蚁蚁巢各品级发生动态,能为当地红火蚁的防控提供技术支撑。【方法】2015年1—12月,在昆明宜良县对红火蚁蚁巢取样、分离,红火蚁取样、称重,统计红火蚁各品级的头数。【结果】地上蚁巢红火蚁蚁群数量发生高峰期在8—9月,其次是3—4月。蚁后数量比例高峰在3—4月;有翅雌蚁比例高峰在4月,其次是7月;有翅雄蚁比例高峰的在5月,其次是9月;生殖蚁幼虫和蛹比例高峰的3、4和12月。地上蚁巢工蚁、兵蚁、职能蚁幼虫和蛹数量发生的高峰期在8—9月,发生小高峰期在4月。【结论】昆明红火蚁蚁群数量动态有2个峰值,第一个峰值为8—9月,以职能蚁防治为主;第二个蜂值3—4月,以生殖蚁和职能蚁防治为主。该结果为昆明红火蚁防控技术提供了数据支撑。     

Biology and oviposition behavior of the phorid Apocephalus attophilus and the response of its host, the leaf-cutting ant Atta laevigata

Atta laevigata (Smith) (Hymenoptera: Formicidae) foragers collected in the field and parasitized by Apocephalus attophilus Borgmeier (Diptera: Phoridae) (1.2% of sampled ants) were larger and had lower survival rates than similar non-parasitized foragers. Moreover, the size of parasitized ants was significantly correlated with the number of A. attophilus puparia and percentage of adult emergence. These results suggest that host size is important for the reproductive success of A. attophilus. Different from most ant-parasitizing phorids, A. attophilus exhibits a pre-oviposition behavior that involves walking towards the host and inspecting it, and this careful approach may be responsible for a relatively low percentage of parasitoid detection by A. laevigata. When an exotic resource (Acalypha sp. leaves) was placed on ants' foraging trails, more foragers were recruited, which resulted in the attraction of A. attophilus. The number and recruiting rates of small workers (minima) were significantly higher on leaves visited by A. attophilus, but larger foragers showed no response to phorids. These results demonstrated that minima react to the presence of A. attophilus and suggest a defensive role of these ants against phorid parasitism.     

Host Location Behavior in a Parasitoid of Imported Fire Ants

Female parasitoids use a hierarchy of cues to locate suitable hosts. We conducted a series of field observations and experiments to examine host location behavior in Pseudacteon tricuspis Borgmeier, a phorid parasitoid of Solenopsis invicta Buren worker ants. The parasitoids were frequently attracted to host workers at disturbed colonies, but were almost never attracted to host workers foraging at baits. When conspecific nonnestmate workers were introduced to baits, resulting in aggressive interactions, parasitoids appeared at the majority of baits. Moreover, larger numbers of parasitoids appeared at baits to which greater numbers of nonnestmate workers had been added. Addition of nonnestmate workers to disturbed colonies resulted in increased numbers of parasitoids attracted. Pseudacteon tricuspis did not display a pattern of uniform distribution at disturbed colonies but often was very abundant at some colony locations while absent or rare at nearby colony locations. Solenopsis invicta workers release alarm pheromones in aggressive interactions with nonnestmates, and this substance is likely an important chemical cue that attracts P. tricuspis flies to host workers from a distance.     

Patterns in foraging and nesting ecology in the neotropical ant, <Emphasis Type="Italic">Gnamptogenys moelleri</Emphasis> (Formicidae,Ponerinae)

Summary This study provides quantitative field data on the natural history and foraging behaviour of the Neotropical bromeliad-nesting ant Gnamptogenys moelleri (Ponerinae) in a sandy plain forest in Southeast Brazil. The ant nested on different bromeliad species and the nests were more frequently found in bigger bromeliads. The species used a wide array of invertebrates in its diet, hunting for live prey and scavenging the majority of the items from dead animals. The food items varied greatly in size (1 to 26 mm). Hunting was always performed by solitary workers. Retrieving was performed by solitary workers (small items), or by a group of 3 to 12 workers recruited to the food source (large items). Almost all G. moelleri foraging activity was restricted to the nest bromeliad. In the warm period more ants left the nest to forage, and foraging trips achieved greater distances compared to the cool season. Trap data revealed that overall availability of arthropod prey is higher in the summer than in the winter. The opportunism in nest site use and in foraging behaviour, the small foraging area, as well as the seasonal differences in foraging activity are discussed and compared with other tropical ants.Received 30 May 2003; revised 22 September 2003; accepted 3 October 2003.     

Efficiency of herbivore exclusion by ants attracted to aphids on the vetch <Emphasis Type="Italic">Vicia angustifolia</Emphasis> L. (Leguminosae)

The efficiency of herbivore exclusion by ants on the vetch Vicia angustifolia L. (Leguminosae) with extrafloral nectary, mediated by ant attraction to aphids was investigated in a field census and laboratory experiments. In the field, workers of Lasius japonicus Santschi and Tetramorium tsushimae Emery frequently visited plants of the vetch parasitized by aphids of Aphis craccivora Koch, but only a few workers visited plants without aphids. An increase in the number of ants visiting a plant with increasing numbers of aphids caused a decrease in the number of larvae of the weevil, Hypera postica Gyllenhal. Therefore, the efficiency of herbivore exclusion by ants was higher on plants parasitized by Ap.craccivora aphids than that on plants unparasitized by aphids. In the laboratory experiments, L.japonicus workers frequently patrolled not only shoots with Ap.craccivora aphids but also shoots without them. However, T.tsushimae workers visited mainly shoots with Ap.craccivora aphids but less frequently on shoots without aphids. Therefore, L.japonicus workers excluded herbivores more efficiently on plants of the vetch than T.tsushimae workers. Consequently, the efficiency of herbivore exclusion by ants on the vetch can be influenced directly by differences in ant species and indirectly by the presence of aphids on plants. The present study highlights the significance of indirect interactions between ants and plants with extrafloral nectary, mediated by ant attraction to aphids for herbivore exclusion of plants.     

Behavioral mechanisms of collective nest-site choice by the ant Temnothorax curvispinosus

This paper examines the individual behavior underlying collective choice among nest sites by the ant Temnothorax (formerly Leptothorax) curvispinosus. Colonies can actively compare options, rejecting a mediocre site when it is paired with a good one, but accepting the same mediocre design if it is instead paired with a worse site. This ability emerges from the behavior of an active minority of workers who organize emigrations. When one of these finds a promising site, she recruits nest mates to it, but only after a delay that varies inversely with site quality. Ants first recruit fellow active ants via slow tandem runs, but eventually switch to speedier transports of the colony’s passive majority. Later transports grow faster still, as ants improve their speed with experience. An ant’s choice of recruitment type is governed by a quorum rule, such that her likelihood of starting to transport increases with the population of the new site. The size of the quorum depends on experience, with ants demanding a larger population to launch immediately into transport than they do to switch to transport after first leading a few tandem runs. Perception of quorum attainment requires direct contact between ants. The ants’ behavior qualitatively matches that of T. albipennis, where models have shown that decentralized choice of the best site depends on quality-dependent recruitment delays, amplified by a quorum rule for initiating transport. Parameter estimates for an agent-based model show significant quantitative differences between the species, and suggest that T. albipennis may place relatively greater emphasis on emigration speed. Received 11 February 2005; revised 10 May 2005; accepted 20 May 2005.     

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

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

Rodents balancing a variety of risks: invasive fire ants and indirect and direct indicators of predation risk

We used foraging trays to compare how oldfield mice, Peromyscus polionotus, altered foraging in response to the presence of fire ants, Solenopsis invicta, and in the presence of direct (predator urine) and indirect (sheltered or exposed microhabitat, moonlight, and precipitation) indicators of predation risk. Foraging reductions elicited by S. invicta were greater than reductions in response to well-documented indicators of risk (i.e., moonlit nights) and the presence of predator urine. The presence of S. invicta always led to reduced foraging, but the overall impact of S. invicta was dependent upon microhabitat and precipitation. When S. invicta was not present, foraging was greater in sheltered microhabitats compared to exposed microhabitats. S. invicta made sheltered microhabitats equivalent to more risky exposed microhabitats, and this effect was especially pronounced on nights without precipitation. The effect of S. invicta suggests that interactions with S. invicta may entail a potentially heavy cost or that presence of S. invicta may represent a more reliable indicator of imminent competition or predation compared to indirect cues of risk and predator urine. The presence of S. invicta led to reduced foraging under situations when foraging activity would otherwise be greatest (i.e., under vegetative cover), potentially reducing habitat quality for P. polionotus and the distribution of seeds consumed by rodents.