Nest Enlargement in Leaf-Cutting Ants: Relocated Brood and Fungus Trigger the Excavation of New Chambers

During colony growth, leaf-cutting ants enlarge their nests by excavating tunnels and chambers housing their fungus gardens and brood. Workers are expected to excavate new nest chambers at locations across the soil profile that offer suitable environmental conditions for brood and fungus rearing. It is an open question whether new chambers are excavated in advance, or will emerge around brood or fungus initially relocated to a suitable site in a previously-excavated tunnel. In the laboratory, we investigated the mechanisms underlying the excavation of new nest chambers in the leaf-cutting ant Acromyrmex lundi. Specifically, we asked whether workers relocate brood and fungus to suitable nest locations, and to what extent the relocated items trigger the excavation of a nest chamber and influence its shape. When brood and fungus were exposed to unfavorable environmental conditions, either low temperatures or low humidity, both were relocated, but ants clearly preferred to relocate the brood first. Workers relocated fungus to places containing brood, demonstrating that subsequent fungus relocation spatially follows the brood deposition. In addition, more ants aggregated at sites containing brood. When presented with a choice between two otherwise identical digging sites, but one containing brood, ants'' excavation activity was higher at this site, and the shape of the excavated cavity was more rounded and chamber-like. The presence of fungus also led to the excavation of rounder shapes, with higher excavation activity at the site that also contained brood. We argue that during colony growth, workers preferentially relocate brood to suitable locations along a tunnel, and that relocated brood spatially guides fungus relocation and leads to increased digging activity around them. We suggest that nest chambers are not excavated in advance, but emerge through a self-organized process resulting from the aggregation of workers and their density-dependent digging behavior around the relocated brood and fungus.     

Oecophylla smaragdina food conversion efficiency: prospects for ant farming

Oecophylla ants are sold at high prices on several commercial markets as a human delicacy, as pet food or as traditional medicine. Currently markets are supplied by ants collected from the wild; however, an increasing interest in ant farming exists as all harvest is easily sold and as ant farming can be combined with the use of the ants in biological control programmes in tropical plantations where pest insects are converted into ant biomass. To assess the cost‐benefits of ant farming based on artificial feeding, food consumption and food conversion efficiency (ECI) of Oecophylla smaragdina (Fabricius) was tested under laboratory conditions. Of the two types of food offered, the ants ingested 76% pure sucrose and 24% insect prey (dry weights) leading to ECI’s of 29% and 39% including brood only or brood plus imago gain, respectively. Based on Thai sugar and protein food costs and ant brood selling prices these efficiencies led to rates of return from 1.52 to 4.56, respectively, if: (i) protein is supplied from commercial products; or (ii) alternatively supplied from free sources such as insects and kitchen waste. These results suggest that Oecophylla ant farming may become highly profitable and deserves further research.     

Fire ant-detecting canines: a complementary method in detecting red imported fire ants

In this investigation, detection dogs are trained and used in identifying red imported fire ants, Solenopsis invicta Buren, and their nests. The methodology could assist in reducing the frequency and scope of chemical treatments for red imported fire ant management and thus reduce labor costs and chemical use as well as improve control and quarantine efficiency. Three dogs previously trained for customs quarantine were retrained to detect the scents of red imported fire ants. After passing tests involving different numbers of live red imported fire ants and three other ant species--Crematogaster rogenhoferi Mayr, Paratrechina longicornis Latreille, and Pheidole megacephala F.--placed in containers, ajoint field survey for red imported fire ant nests by detection dogs and bait traps was conducted to demonstrate their use as a supplement to conventional detection methods. The most significant findings in this report are (1) with 10 or more red imported fire ants in scent containers, the dogs had >98% chance in tracing the red imported fire ant. Upon the introduction of other ant species, the dogs still achieved on average, a 93% correct red imported fire ant indication rate. Moreover, the dogs demonstrated great competence in pinpointing emerging and smaller red imported fire ant nests in red imported fire ant-infested areas that had been previously confirmed by bait trap stations. (2) Along with the bait trap method, we also discovered that approximately 90% of red imported fire ants foraged within a distance of 14 m away from their nests. The results prove detection dogs to be most effective for red imported fire ant control in areas that have been previously treated with pesticides and therefore containing a low density of remaining red imported fire ant nests. Furthermore, as a complement to other red imported fire ant monitoring methods, this strategy will significantly increase the efficacy of red imported fire ant control in cases of individual mount treatment.     

Evidence of intracolony transmission of Thelohania solenopsae (Microsporidia: Thelohaniidae) in red imported fire ants (Hymenoptera: Formicidae) and the first report of spores from pupae.

Red imported fire ant, Solenopsis invicta, colonies were infected horizontally by introducing live brood (mainly larvae and pupae) infected with Thelohania solenopsae. Live, infected brood introduced into uninfected colonies were adopted and raised to adulthood instead of being executed by the recipient colony. Introductions of infected larvae with uninfected pupae, which eclose into adult worker caste fire ants, resulted in an 80% infection rate of the inoculated colonies. Infections from introductions of infected pupae with uninfected larvae resulted in a 37.5% infection of inoculated colonies. Infections were also detected in 11.6 and 3.7% of the adult worker caste ants that eclosed from uninfected large larvae and pupae, respectively, that were held with infected adult workers. Microscopic examination of infected brood revealed sporoblasts and large numbers of spores of T. solenopsae in S. invicta pupae.     

The economics of brood raiding and nest consolidation during ant colony founding

Summary The most dangerous time for an ant colony is during the founding stage when the small colony is vulnerable to predation and competition. Colonies can grow more rapidly when multiple queens cooperate in raising the first worker brood (pleometrosis) or by raiding other incipient colonies for their brood. This brood raiding has been proposed to be the primary force selecting for pleometrosis, i.e. multiple-queen colonies may have a considerable advantage in destroying neighbours by aggressively stealing their brood. An alternative hypothesis is that incipient nests are part of a larger, interconnected population structure and that brood raiding reflects cooperative pleometrosis with subdivided colonies. A simple mathematical model supports the second hypothesis: workers of incipient colonies are especially favoured to peaceably abandon their nest and join with other colonies if the queens are related or queens from raided colonies can infiltrate the raiding colony. The latter condition is often met in ant species that brood raid and particularly exemplified in fire ants (Solenopsis invicta), where brood raiding involves little mortal combat and combines with pleometrosis to rapidly increase colony size. It is proposed that the term nest consolidation should replace brood raiding to more accurately reflect the relatively non-aggressive and potentially apparently cooperative nature of interactions between incipient ant colonies.     

Larger laboratory colonies consume proportionally less energy and have lower per capita brood production in Temnothorax ants

Colony size can affect individual- and colony-level behavioral and physiological traits in social insects. Changes in behavior and physiology in response to colony growth and development can affect productivity and fitness. Here, we used respirometry to study the relationship between colony size and colony energy consumption in Temnothorax rugatulus ants. In addition, we examined the relationship between colony size and worker productivity measured as per capita brood production. We found that colony metabolic rate scales with colony size to the 0.78 power and the number of brood scales with the number of workers to the 0.49 power. These regression analyses reveal that larger ant colonies use proportionally less energy and produce fewer brood per worker. Our findings provide new information on the relationships between colony size and energetic efficiency and productivity in a model ant genus. We discuss the potential mechanisms giving rise to allometric scaling of metabolic rate in ant colonies and the influence of colony size on energy consumption and productivity in general.     

Capital and income breeding and the evolution of colony founding strategies in ants

Ability to store resources that will be used for reproduction represents a potential life history adaptation because storage permits feeding and reproduction to be decoupled spatially and/or temporally. The two ends of a continuum involve acquiring all resources prior to reproduction (capital breeding) or acquiring all resources during the reproductive period (income breeding). Traditional life history theory examines tradeoffs between costs and benefits of such strategies, but this theory has not been integrated into life history studies of ants, even though founding queens have the analogous strategies of fully claustral (capital breeding) and semi-claustral (income breeding). This study demonstrates that facultatively semi-claustral queens of the seed-harvester ant Pogonomyrmex desertorum exhibit phenotypic plasticity during colony founding because unfed queens produced few, small minims, whereas ad libitum fed queens produced larger, heavier minims and additional brood. Fed queens also lost less mass than unfed queens despite their producing more brood. Overall, foraging provides queens with a suite of benefits that likely offset potential negative effects of foraging risk. Life history studies across a diverse array of taxa show that capital breeding is consistently associated with low availability and/or unpredictability of food, i.e., environmental conditions that favor prepackaging of reproductive resources. Such a broad and consistent pattern suggests that similar factors favored the evolution of fully claustral (capital breeding) colony founding in ants. Overall, these data suggest that ant researchers should revise their conventional view that fully claustral colony founding evolved because it eliminated the need for queens to leave the nest to forage. Instead, colony founding strategies should be examined from the perspective of environmental variation, i.e., availability and predictability of food. I also provide a functional scenario that could explain the evolution of colony founding strategies in ants. Received 16 November 2005; revised 1 March 2006; accepted 29 March 2006.     

Susceptibility and behavioral response of red imported fire ant (Hymenoptera: Formicidae) to selected entomogenous nematodes (Rhabditida: Steinernematidae & Heterorhabditidae).

Pathogenicity of infective juveniles of selected Steinernema spp. and Heterorhabditis spp. toward developing and reproductive stages of the red imported fire ant, Solenopsis invicta Buren, was tested under laboratory conditions. At 10(3)-10(5) infective juveniles per Petri dish, mortality of reproductive larvae, pupae, and alates ranged from 28 to 100% at higher doses after 96 h at 23-25 degrees C. Steinernema carpocapsae All was the most consistent species tested; this nematode caused mortality of fire ant larvae, pupae, and alates of 82-94, 64-96, and 38-99%, respectively. Although not susceptible to nematode infection, worker ants vigorously preened nematodes from brood, alates, and themselves. In a field study, S. carpocapsae (5 x 10(6) and 2 x 10(6) drench, 2 x 10(6) infective juvenile infection) was applied to active fire ant mounds in 3.8-liter suspensions. Hydramethylnon (75 ml), a water drench, a water injection, and untreated fire ant mounds were marked and treated. Overall activity in mounds treated with nematodes of hydramethylnon ranged from 40 to 48%. Satellite mound activity accounted for 32-44% of overall activity in mounds treated with nematodes 2 wk after treatment. However, 6 wk after treatment, activity in mounds treated with hydramethylnon was 44%; activity of mounds treated with nematodes ranged from 52 to 80%. Satellite mound activity accounted for 0-24% of overall activity. Whereas a soil drench of S. carpocapsae showed potential as a control method for the red imported fire ant, colony relocation after nematode treatment could limit overall efficacy unless application techniques are developed to overcome or take advantage of the movement.     

Thermoregulatory brood transport in the fire ant, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>

Nest structure in ants is often designed to optimize the colony’s ability to thermoregulate, and this specialization is most highly developed in mound-building ant species. Solenopsis invicta invest a large amount of energy in building mounds and transporting their brood up and down in their nests as a means of thermoregulation. Because few ant species build true mounds, we wanted to determine the effectiveness of these mounds in harvesting solar heat as well as to distinguish what factors (temperature vs. circadian rhythm) govern where fire ants place their brood in the mound and when they place it. We measured temperature patterns in the mound over several days at different depths and under different conditions (under direct sunlight or shade), and then conducted a series of field experiments to manipulate the orientation and time of heating. On cool mornings in spring or fall, surface temperatures of the mound rise at the fastest rate on the side receiving the most direct sunlight (usually the south side). This heating causes a temperature gradient through different depths in the mound, and shows little difference from outside ground temperature at a depth greater than ~40 cm inside the nest. In the morning, fire ants move their brood up into the mound on the side most directly heated, and when temperatures exceed optimal (~32°C) they move their brood down the temperature gradient to lower depths in the nest. In addition to this, mound temperature does not only increase due to direct sunlight, but temperature also increases higher than ground temperatures when the mound is in the shade due to its low specific heat. Experiments in which sunlight was mirrored to the normally shaded side of the mound, or when mounds were heated at night, revealed that S. invicta primarily track temperature patterns and do not rely on behavioral habits or circadian rhythms for the thermoregulatory transport of their brood. When mounds were shaded, S. invicta brood was evenly distributed directly under the surface of the mound rather than aggregating towards a specific side. The fire ant mound is important for thermoregulation because, compared to moundless subterranean nests, it absorbs heat more rapidly both in direct sunlight and shady conditions. Temperature tracking within the nest is key to understanding thermoregulatory placement of fire ant brood, as well as insight into the production of sexual brood and reproduction. Received 9 August 2007; revised 31 January 2008; accepted 7 February 2008.     

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.     

Intraspecific usurpation of incipient fire ant colonies

Brood raiding, or reciprocal stealing of brood, is common amongincipient colonies of the fire ant Solenopsis invicta. Paradoxically,workers from a colony that loses its brood during a raid oftenabandon their nest and join the winning colony. Queens abandonedby their workers may then migrate from their original nest siteand attempt to forcefully usurp another incipient colony bydisplacing that colony's queen or queens. This study examinedfactors that influence the success of usurpation attempts. Queensattempting to usurp a nearby colony after laboratory brood raidswere successful in less than 30% of trials. Usurpation attemptswere more successful if workers familiar to the migrating queenwere present in the nest, as would happen if a queen were tofind the colony to which her workers had moved. Cross-fosteringexperiments showed that this effect was due to familiarity ratherthan relatedness. Usurpation attempts were less successful ifthey were delayed by 16 h. The probability of usurpation successwas not reduced by doubling the number of defending workersin the invaded colony. However, colonies founded by three queenswere almost always able to resist usurpation attempts. Theseresults support the hypothesis that workers abandon their natalcolony after losing a brood raid to increase the likelihoodthat their queen can usurp the colony to which they migrate.These results also provide the first evidence that coloniesof ants founded by several queens are better able to resistusurpation attempts than colonies founded by a single queen     

Coupled adult-brood transport augments relocation in the Indian queenless ant Diacamma indicum

Colony relocation is an important aspect in the lives of social insects. In ants, the process of relocation is further complicated as brood, in addition to adults, have to be transported to the new nest. Here, we have investigated brood transport in the Indian ponerine ant Diacamma indicum, which uses tandem running—a primitive mode of recruitment—for the entire colony to relocate. We have found that there were no brood transport specialists and most of the brood was transported in the mandibles of followers that were being tandem run. Therefore, in a single tandem run, one adult and one brood item was effectively transported by tandem leaders augmenting the relocation process.     

Aggressive interactions between Solenopsis invicta and Linepithema humile (Hymenoptera: Formicidae) under laboratory conditions

The Argentine ant, Linepithema humile (Mayr), and the red imported fire ant, Solenopsis invicta Buren, are natural agonists in their country of origin. Since the first report of L. humile in California in 1907 its range expanded statewide, displacing native ant species wherever it spread. Since the discovery of established populations of S. invicta in southern California in 1998, it has been restricted to discrete areas of southern California. However, as these discrete populations expand, they are encountering large populations of L. humile, which are effective competitors for available resources and are particularly aggressive in their encounters with other ant species such as S. invicta. Most Dolichoderine ants such as L. humile do not prefer to forage on baits made with defatted corn grit and soybean oil typically used in red imported fire ant control programs. Applications of these baits in areas where distributions of these species overlap might selectively affect populations of S. invicta and give L. humile a competitive advantage. Three laboratory experiments were conducted to determine the competitive outcomes between S. invicta pitted against L. humile: 1) agonistic behavior of workers in small arenas, 2) colony interactions with different population ratios, and 3) the effects of pyriproxyfen on the competitiveness of S. invicta against L. humile. Populations of S. invicta consisting of major workers killed more L. humile than did minors or a mixture of majors and minors. When paired against L. humile colonies consisting of 1,100 workers, colonies consisting of 38 S. invicta workers were easily defeated by L. humile. Colonies consisting of 450 S. invicta workers plugged their nest entrances, but they were ultimately defeated by L. humile after 13 d. The S. invicta colonies consisting of 1,100 workers took control of the bridge connecting the colonies, invaded the L. humile colony, killed the Argentine ant queens, and removed their brood. Pyriproxyfen-treated fire ant workers took significantly longer to chase the Argentine ants from a connecting bridge than did untreated fire ants. Thus, fire ant baits may have long-term effects on intercolonial aggression between S. invicta and L. humile, especially when Argentine ant populations are high in the summer.     

The cost of being queen: Investment across Pogonomyrmex harvester ant gynes that differ in degree of claustrality

The role of the ant colony largely consists of non-reproductive tasks, such as foraging, tending brood, and defense. However, workers are vitally linked to reproduction through their provisioning of sexual offspring, which are produced annually to mate and initiate new colonies. Gynes (future queens) have size-associated variation in colony founding strategy (claustrality), with each strategy requiring different energetic investments from their natal colony. We compared the per capita production cost required for semi-claustral, facultative, and claustral gynes across four species of Pogonomyrmex harvester ants. We found that the claustral founding strategy is markedly expensive, costing approximately 70% more energy than that of the semi-claustral strategy. Relative to males, claustral gynes also had the largest differential investment and smallest size variation. We applied these investment costs to a model by Brown and Bonhoeffer (2003) that predicts founding strategy based on investment cost and foraging survivorship. The model predicts that non-claustral foundresses must survive the foraging period with a probability of 30–36% in order for a foraging strategy to be selectively favored. These results highlight the importance of incorporating resource investment at the colony level when investigating the evolution of colony founding strategies in ants.     

Assessing the effects of sodium on fire ant foraging in the field and colony growth in the laboratory

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The biogeography of prediction error: why does the introduced range of the fire ant over‐predict its native range?

Aim The use of species distribution models (SDMs) to predict biological invasions is a rapidly developing area of ecology. However, most studies investigating SDMs typically ignore prediction errors and instead focus on regions where native distributions correctly predict invaded ranges. We investigated the ecological significance of prediction errors using reciprocal comparisons between the predicted invaded and native range of the red imported fire ant (Solenopsis invicta) (hereafter called the fire ant). We questioned whether fire ants occupy similar environments in their native and introduced range, how the environments that fire ants occupy in their introduced range changed through time relative to their native range, and where fire ant propagules are likely to have originated. Location We developed models for South America and the conterminous United States (US) of America. Methods We developed models using the Genetic Algorithm for Rule‐set Prediction (GARP) and 12 environmental layers. Occurrence data from the native range in South America were used to predict the introduced range in the US and vice versa. Further, time‐series data recording the invasion of fire ants in the US were used to predict the native range. Results Native range occurrences under‐predicted the invasive potential of fire ants, whereas occurrence data from the US over‐predicted the southern boundary of the native range. Secondly, introduced fire ants initially established in environments similar to those in their native range, but subsequently invaded harsher environments. Time‐series data suggest that fire ant propagules originated near the southern limit of their native range. Conclusions Our findings suggest that fire ants from a peripheral native population established in an environment similar to their native environment, and then ultimately expanded into environments in which they are not found in their native range. We argue that reciprocal comparisons between predicted native and invaded ranges will facilitate a better understanding of the biogeography of invasive and native species and of the role of SDMs in predicting future distributions.     

Dynamics of colony emigration in the ant Aphaenogaster senilis

In many social insect species, colonies frequently emigrate to a new nest. This requires the coordination of many individuals, and it puts the queen at risks of being lost or predated. We experimentally studied colony emigration in the ant Aphaenogaster senilis, who emigrates frequently and obligatorily reproduces by colony fission. As in other species, colony emigration was characterised by a synchronised relocation of workers. Foragers found the new nest site and triggered the relocation of the “inside” workers, which built up following a sigmoid curve. Unlike in Temnothorax, where workers are transported to the new nest, most individuals relocated by walking. The brood was transported around the middle of colony relocation, mostly by “inside” workers because they represent most of the workforce. The queen walked to the new nest at the middle of colony relocation, when the flow of ants to the new nest was maximal. Overall, this temporal dynamic of colony emigration is similar to that observed in other species. However, we argue that species-specific traits, such as whether workers are transported to the new nest or relocate by themselves, may affect parts of the process of colony emigration.     

Effects of colony-level attributes on larval feeding in the fire ant, Solenopsis invicta

Summary: Do colony attributes modulate individual behavior? The effects of colony size and worker:brood ratio on the rate of worker-to-larva trophallaxis in the fire ant, Solenopsis invicta, were investigated. Neither colony size ranging from 100 to 10,000 nor worker:brood ratio ranging from 1:1 to 16:1 affected the density of workers on the brood pile, nor the rate or duration of worker-to-larva trophallaxis. The demands of hungry larvae were met even in groups as small as 100 workers in worker:brood ratios as small as 1. Only when the worker:brood ratio was less than 1, were larvae tended or fed at reduced rates. Under natural conditions, this occurs only in incipient colonies. Otherwise, in post-incipient colonies, the flow of food to larvae was unmodified by colony attributes. The implications of this finding are two-fold: First, it reinforces previous research demonstrating that social feeding in the fire ant emerges from localized interactions rather than mass communication. Second, it highlights the resiliency of this weedy species. Hypothetically, colonies drastically reduced by catastrophic events such as flooding should still be able to produce sexuals.     

Experimental evidence for weak effects of fire ants in a naturally invaded pine‐savanna ecosystem in north Florida

1. Fire ants naturally invade some undisturbed ecosystems of high conservation value and may negatively impact co‐occurring ants. 2. Over 3 years, fire ants were added and removed from a longleaf pine savanna ecosystem that naturally supports a low density of fire ants. Impacts on co‐occurring ants were monitored using pitfall traps. 3. Treatments resulted in significant differences in average fire ant abundance across all plots only in the first year of the experiment. Fire ants had little discernible impact. The abundance and species richness of co‐occurring ants in removal plots never differed from unmanipulated control plots. The abundance of co‐occurring ants was very slightly lower and ant species richness was slightly higher where Solenopsis invicta Buren colonies were added, but neither contrast was significant. 4. The poor conditions in this habitat for many native ants may explain this outcome. More broadly, the impact of fire ants on ant assemblages still appears to be secondary and largely a consequence of human impacts on the environment.