Queen recruitment in a multiple-queen population of the fire ant Solenopsis invicta

We assessed patterns of new queen recruitment in a polygyne(multiple queens per nest) population of the fire ant Solenopsisinvicta in its introduced range. Newly recruited queens wereidentified using four physiological markers, and genotypic datafrom nuclear and mitochondrial markers were used to estimaterelatedness of new nest mate queens to each other and to theolder nest mate queens. In total, 1.7% of the queens collectedin late spring and early summer were deemed to be new recruits.The relatedness of these queens to each other and to the olderqueens within nests was not significantly different from zero,suggesting that newly recruited queens represent a random sampleof potential reproductive queens in the population. Moreover,the number of new queens recruited within nests was not correlatedwith the number of older queens present and did not differ significantlyfrom a Poisson distribution. Thus, queen recruitment in this populationof S. invicta appears to occur at random with respect to thenumber of older queens present within nests.     

Reproductive development and ontogeny of queen pheromone production in the fire ant Solenopsis invicta

Queens of the fire ant, Solenopsis invicta Buren, produce several releaser and primer pheromones. Using bioassays, the ontogeny of three of these pheromones related to reproductive development was investigated. Virgin queens, in which the process of wing-shedding (dealation) serves as an indicator of the initiation of reproductive development, were studied. First, the production of two queen pheromones produced in the poison gland was examined. The pheromone responsible for initiation and maintenance of retinue formation, a releaser effect, was found to be produced in detectable quantities 2 days after dealation, at which time queens showed significant ovary development and many (30%) had started laying eggs. A primer pheromone that inhibits alate virgin queens from dealating was detected in queens 3 days following wing-shedding, when 80% of the queens were ovipositing. Second, I examined the onset of a pheromone of unknown glandular origin produced by reproductively active virgin queens which leads to their destruction in queenright colonies by stimulating workers to attack and kill them. This pheromone is secreted in quantities detectable by bioassay 2 days after dealation. Thus, in S. invicta, the ontogeny of three distinct queen pheromones is tightly linked with ovary development and initiation of egg laying. These results demonstrate reproductive and communicative functions are closely associated during the transition from potential to functional queen.     

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

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

A formal assessment of gene flow and selection in the fire ant Solenopsis invicta

Abstract.— Recent studies of the introduced fire ant Solenopsis invicta suggest that introduced polygyne (with multiple queens per nest) populations are strongly influenced by male-mediated gene flow from neighboring monogyne (single queen per nest) populations and selection acting on a single locus, general protein-9 (Gp-9) . This investigation formally tests this hypothesis and determines if these processes can account for the genotypic structure of polygyne S. invicta . To increase the statistical power of this test, we considered the genotypes of polygyne queens and workers at both Gp-9 and the closely linked, selectively neutral locus Pgm-3 . We then constructed and analyzed a novel mathematical model to delimit the effects of monogyne male gene flow and selection on the joint genotypes at the Pgm-3/Gp-9 superlocus. Using this framework, a hierarchical maximum-likelihood method was developed to estimate the best-fitting gene flow and selection parameters based on the fit of our model to data from both the current study and an earlier one of the same population. In each case, selection on polygyne queens and workers alone, with no monogyne male gene flow, provides the most parsimonious explanation for the observed genotype frequencies. The apparent discrepancy between this result and the empirical evidence for monogyne male gene flow indicates that undocumented factors, such as other forms of selection in polygyne males or workers, are operating in introduced polygyne S. invicta .     

The red imported fire ant,Solenopsis invicta,modifies predation at the soil surface and in cotton foliage

Red imported fire ants, Solenopsis invicta, are generalist predators that can have major impacts on foliar arthropod communities in agricultural systems; however, their effects as predators at the soil surface have not been adequately characterized. We examined the contribution of fire ants to predation at the soil surface and in cotton foliage at two sites and over the course of two field seasons in Georgia, using egg masses of the beet armyworm, Spodoptera exigua. To assess interactions between fire ants and other arthropod species, we also measured the densities of edaphic predators and honeydew‐producing hemipterans at both sites. The sites occurred in different growing regions (Piedmont and Coastal Plain), and allowed us to characterize the importance of fire ants as predators under different climatic and soil conditions. Fire ant suppression decreased egg predation at both field sites, and predation by fire ants at the soil surface was equal to if not greater than that in cotton foliage. However, the impact of fire ants on predation varied between sites, likely due to differences in climate and the composition and activity of the extant arthropod communities. Our study also indicates that fire ant suppression is associated with decreases in the density of honeydew‐producing insects, and increasing abundance of whiteflies on the plants coincided with a decrease in egg predation at the soil surface. This finding suggests the mutualism between ants and whiteflies may lead to a shift in predation intensity from edaphic towards plant‐based food webs.     

几种饵料对红火蚁觅食的引诱作用

在荒地类型生境中测定花生酱、猫饲料、狗饲料、花生油、面包及蜂蜜等几种常见饵料对红火蚁的引诱作用。结果表明:面包、狗饲料、猫饲料和花生酱对红火蚁引诱力较强,花生油次之,蜂蜜最低;选择性试验结果表明红火蚁对供试的几种饵料无明显选择性。     

红火蚁的社会型及其相关基因Gp-9的研究综述

红火蚁Solenopsis invicta Buren的蚁巢具有2种基本的社会组织形态:单后型和多后型。蚁巢中蚁后的数量是由工蚁和蚁后的基因型共同决定的。红火蚁的Gp-9基因编码一种气味结合蛋白并控制这种社会多态性的表达。Gp-9基因的发现首次证明单个基因在昆虫复杂的社会行为中起关键的作用。从红火蚁的社会型特征、Gp-9基因与不同社会型的关系以及Gp-9基因的相关研究对该领域的研究进展进行综述。     

Effect of queen number on the production of sexuals in natural populations of the fire ant, Solenopsis invicta

ABSTRACT. To determine whether the production of sexuals in the fire ant, Solenopsis invicta Buren, is related to queen number in the field, the numbers of sexuals in monogyne (M) and polygyne (P) colonies of this species were compared. Large colonies ( n =25 M and 25 P) were collected at random during spring, summer and fall when sexuals are actively produced, and the numbers of sexuals present were counted. The numbers of alates plus sexual pupae and larvae in M colonies far exceeded those in P colonies in each season. Significant differences between M and P colonies were also found when the numbers of alates and sexual immatures (pupae + larvae) were compared separately in each season. In addition, M colonies contained significantly more female sexuals (alates + pupae) than did P colonies in the summer and fall, and significantly more males in all three seasons. The negative relationship between queen number and number of sexuals provides evidence that queen control over the production of sexuals, previously established in laboratory experiments, also occurs under natural conditions.     

研究红火蚁工蚁活动性的新方法-刻度图示法

应用一种新方法--刻度图示法研究红火蚁工蚁Solenopsis invicta Buren群体活动性规律,结果表明,标尺上红火蚁最高爬升高度、平均爬升高度、工蚁恢复平静时间、工蚁累计数量均随着蚁巢体积增大而增大,通过分析建立红火蚁工蚁活动性程度各指标值与蚁巢大小关系多个模型.     

Trophallaxis by temporal subcastes in the fire ant, Solenopsis invicta, in response to honey

ABSTRACT. Trophallactic behaviour of temporal subcastes in Solenopsis invicta Buren colonies was studied in two separate experiments. First, radiolabeled honey was presented to small colonies containing larvae, queens and marked workers; radioactivity in the head, thorax and abdomen of each ant was measured after 0.25, 1, 6, 12, 24, 48 and 72 h. The quantity of honey per subcaste was dependent on the total amount of honey collected. As more honey entered the colony, the quantity in foragers, queens and reserves increased more rapidly than that in nurses or larvae. The internal distribution of radioactivity in the head, thorax or abdomen indicated that differences between subcaste members in either the rate of food exchange or digestion existed. In a second experiment behavioural observations were made on marked workers in colonies before and after the presentation of honey. Reserves spent significantly more time engaged in trophallaxis than either foragers or nurses, both as active donors and active receivers. Based on number of encounters, nurses were more likely to receive honey while reserves and foragers were more likely to donate honey. The duration of encounters involving nurses tended to be shorter than those involving reserves or foragers.     

The success of alternative reproductive tactics in monogyne populations of the ant Solenopsis invicta: significance for transitions in social organization

Newly produced queens from monogyne (single-queen) coloniesof the ant Solenopsis invicta usually initiate reproductionindependently, that is, without worker assistance. However,some recently mated queens attempt to bypass this risky phaseof new colony foundation by entering established nests to reproduce,although it is unclear how often these queens are successfulin natural populations. We surveyed a mature monogyne populationof S. invicta in both 1995 and 1996 for colonies headed by queensincapable of independent colony founding (diploid-male-producingqueens) in order to estimate the frequency of colonies thatare headed by queens that initiated reproduction within establishednests (adopted queens). Using the frequency of diploid-male-producingqueens among the recently mated queens in this population, weestimated that the overall rate of queen replacement by adoptedqueens is about 0.7% per colony per year. Although theory suggeststhat a change to a novel queen reproductive tactic could beassociated with a fundamental change in social organization(queen number), this does not appear to be the case in monogyneS. invicta. However, the evolution of nest-infiltrating reproductivetactics by queens in a monogyne population and the evolutionof multiple-queen societies may result from similar ecologicalpressures facing newly mated queens. We therefore incorporatethis strategy into an existing theoretical framework that wasdeveloped to explain the evolution of alternative social organizationsin ants, providing testable predictions regarding the distributionand frequency of queen adoption in other single-queen ant societies.     

Characteristics of 11 polymorphic microsatellite markers in the red imported fire ant, Solenopsis invicta Buren

We have characterized 11 polymorphic microsatellite loci in the invasive ant Solenopsis invicta. Primer pairs were evaluated on fire ants collected from monogyne mounds in Lauderdale County, Mississippi. The observed and effective number of alleles ranged from two to six and from 1.31 to 2.64, respectively. The observed and expected heterozygosity values ranged from 0.1613 to 0.7826 and from 0.1491 to 0.6242, respectively. The polymorphism information content of the microsatellites ranged from 0.1482 to 0.6208. Probability tests indicated significant deviations from the Hardy-Weinberg equilibrium at three loci. Pairwise tests did not detect linkage disequilibrium between any pair of loci.     

Behaviour of worker subcastes in the fire ant, Solenopsis invicta, in response to proteinaceous food

ABSTRACT. The effects of division of labour on response behaviour to food in the red imported fire ant, Solenopsis invicta Buren, were examined to determine if caste members differ in amount of food taken, in rate of food transfer, or in internal distribution of food; and to see if food availability, time, or temporal subcaste pairing affect feeding behaviour. To measure differences in behaviour we fed radioiodinated albumin mixed with egg yolk to colonies containing larvae, queens, and (a) foragers and nurses, or (b) foragers and reserves, or (c) nurses and reserves. Samples were taken over a 72-h period and radioactivity in the head, thorax and abdomen of each worker was determined. There were significant differences between nurses, foragers and reserves in quantity of food consumed, rate of transfer, and internal distribution of radioactivity. These differences were related to their respective roles of foraging, food storage and transfer, and brood tending. The quantity of food taken per subcaste was dependent on the total amount of food in the colony, with transfer rates differing between subcastes as the quantity of food in the colony increased. The rate at which protein was transferred between subcastes was slower in the reserves than that in either foragers or nurses. Therefore, reserves may serve as a temporary store of protein for the colony.     

研究红火蚁工蚁活动性的新方法—刻度图示法

应用一种新方法——刻度图示法研究红火蚁工蚁Solenopsis invicta Buren群体活动性规律,结果表明,标尺上红火蚁最高爬升高度、平均爬升高度、工蚁恢复平静时间、工蚁累计数量均随着蚁巢体积增大而增大,通过分析建立红火蚁工蚁活动性程度各指标值与蚁巢大小关系多个模型。     

Successful establishment of the invasive fire ant Solenopsis invicta in Taiwan: insights into interactions of alternate social forms

Aim Understanding the factors underlying the successful establishment of invasive ant species is critical for developing quarantine strategies to prevent additional invasions as well as for determining how such species overcome the selective pressures in invaded areas. Although several studies have revealed differences in the social organization and population genetics of invasive ants in their native and introduced ranges, few studies have considered the potential interactions between alternate social forms within newly‐invaded areas simply because many invasive ants are characterized as polygyne or unicolonial. Location Taoyuan and Chiayi County, Taiwan. Methods Both social forms (polygyne and monogyne) of the red imported fire ant, Solenopsis invicta, occur in two separately invaded areas in Taiwan (Taoyuan and Chiayi). We employed intensive sampling methods and diagnostic polymerase chain reaction assays to determine the distribution of these two social forms in both infested areas in Taiwan. Results The distribution of social forms differs dramatically between the two infested areas, consistent with different invasion histories regarding the original make‐up of individuals comprising the initial founder group. The Taoyuan population likely was colonized initially by ants of both social forms, with the subsequent spread characterized by continuous outward movement of the two forms, particularly the monogyne form. In contrast, the initial founders of the Chiayi population likely were of the polygyne social form only, suggesting the monogyne social form in this population appeared only recently and likely arose directly from polygyne queens converting to the alternate social form. Main conclusions Our results provide detailed insights into the invasion history of S. invicta in Taiwan and suggest that the distinct reproductive biology of the two social forms may have shaped the current distribution of S. invicta in these infested areas and that the dynamics of two forms may affect the long‐term persistence and potential for spread of this pest ant species.     

Group defense by colony-founding queens in the fire ant Solenopsis invicta

Mutualistic associations among nonkin can form when animalsin groups have a greater chance of overcoming challenges presentedby the environment than do solitary animals. Colony foundingby small groups of unrelated queens, a habit documented in severalspecies of ants, is often interpreted as a mutualistic interactionselected by intense competition among incipient colonies. However,many new colonies in these species are founded in areas wheretheir chief enemies are mature ant colonies, rather than othernewly founded colonies. In this study, we tested whether groupnest-founding in the fire ant Solenopsis invicta improved theability of queens to survive attacks by mature colonies. Inthe laboratory, queens in groups of three were more likely thansolitary queens to survive attacks by workers of the nativefire ant Solenopsis geminata. When newly mated queens were establishedexperimentally in the field, workers from mature S. invictacolonies caused the majority of queen deaths. Queens in groupsof two, but not in groups of four, had higher survival ratesthan did solitary queens during the period between colony establishmentand the appearance of the first workers. The advantage of cooperativedefense approximately counterbalanced the disadvantages causedby competition within foundress associations of two to threequeens. Previous studies have shown that colonies founded bymultiple queens produce larger worker populations than coloniesfounded by solitary queens; however, experimentally increasingworker number in incipient colonies had no effect on colonysurvival in the field.     

不同方法对红火蚁社会型鉴定效果的比较

红火蚁Solenopsis invicta Buren的社会型——单蚁后(Monogyne)和多蚁后(Polygyne)型的鉴定方法众多,对于鉴定方法之间的比较较少。为了探究鉴定红火蚁社会型的有效实验方案,本研究对采集自中国南方的红火蚁种群样品进行了生物学观察,并采用了几种常用的方法对样品社会型进行了测定,此外还对比了4种不同DNA聚合酶在实验中的效果。多重PCR技术(Multiplex Polymerase Chain Reaction)、Gp-9 b等位基因扩增两种方法成功率高,能准确鉴定红火蚁社会型。4对同时扩增B、b等位基因的引物成功率不高。实验结果表明,将多重PCR技术、Gp-9 b等位基因扩增,与生物学观察相结合,能更准确地对红火蚁社会型进行鉴定。在PCR实验中选用具有适度保真性和热启动效果的DNA聚合酶能提高红火蚁社会型鉴定的准确性。     

On the relationship between queen number and fecundity in polygyne colonies of the fire ant Solenopsis invicta

ABSTRACT. The reproductive status of individual queens in relation to the number of queens in polygyne colonies of Solenopsis invicta Buren was investigated under both field and laboratory conditions. In field colonies, the weight of individual queens is inversely related to the number of resident queens present. The weight of a queen is positively correlated with fecundity, as measured by the number of eggs laid in 5 h in isolation. Thus increasing queen number has a strong negative effect on the fecundity of individual queens in these colonies. A similar relationship was established in standardized laboratory colonies using queens from a single experimental population. Colony size by itself had no significant effect on queen weight in either the field or laboratory colonies, suggesting that the inverse relationship between queen number and fecundity results from mutual inhibition among queens, possibly involving pheromones, rather than reduced nutrition through lower worker/queen ratios. While uninseminated queens can comprise up to 50% of the functional (i.e. egglaying) queens in polygyne colonies of this ant, they tend to be less fecund than their inseminated nestmates; uninseminated queens within a given colony generally weigh 5–25% less.