Jules Silverman

You are what you eat: diet modifies cuticular hydrocarbons and nestmate recognition in the Argentine ant, Linepithema humile.

Abstract Nestmate recognition plays a key role in the behavior and evolution of social insects. We demonstrated that hydrocarbons are the chemical cues used in Argentine ant, Linepithema humile, nestmate recognition, and that these hydrocarbons can be acquired from insect prey. Consequently, Argentine ant cuticular hydrocarbon patterns reveal the same hydrocarbons present in their diet. Diet alters both the recognition cues present on the cuticular surface and the response of nestmates to this new colony odor, resulting in aggression between former nestmates reared on different insect prey.

Nestmate discrimination in ants: effect of bioassay on aggressive behavior

Summary: Aggression assays are commonly used to study nestmate recognition in social insects. Methods range from detailed behavioral observations on small numbers of insects to counts of individuals fighting in group interactions. These assays vary in the equipment used and the intensity and duration of observations. We used the Argentine ant, Linepithema humile, to compare four aggression bioassays for consistency between replicates, similarity between assays, and ability to predict whole colony interactions. The assays included were 1 live – 1 dead ant interactions, live 1-1 battles, live 5-5 battles, and 1 ant introduced to a foreign colony. We tested six ant colonies in all pairwise combinations using four different assays and two to three scoring methods per assay. We also conducted a colony merging experiment to see which assays were capable of predicting this ecologically important event. We found that scoring methods within assays yielded very similar results, giving us no reason to favor observationally intense procedures, such as continuous scanning, over less observationally intense systems, such as snapshot surveys. Assays differed greatly in their consistency between replicates. No two replicates of the 1 live – 1 dead assay were significantly correlated. The live 5-5 and the colony introduction assays were the most consistent across replicates. The mean scores of the live 1-1, live 5-5 and colony introduction assays were all significantly correlated with each other; only the live 5-5 assay was significantly correlated with the 1 live – 1 dead assay. Assays that utilized the greatest number of live ants were the most likely to reveal high levels of aggression. The aggression scores of all but the 1 live – 1 dead assay were positively correlated with the number of ants that died during whole colony encounters and negatively associated with colony merging. We conclude that all live ant assays tested are useful tools for analyzing aggressive interactions between colonies, but that the pairing of a live and dead ant produced inconsistent results and generally lower levels of aggression. We found relatively low consistency between trials using the live 1-1 assay, but found that with sufficient replication its results were highly correlated with the assays using more interacting ants. We suggest that isolated aggressive acts in assays do not necessarily predict whole colony interactions : some colonies that fought in bioassays merged when the entire colonies were allowed to interact.

Glucose aversion in the german cockroach, Blattella germanica

Abstract A recently discovered strain of German cockroaches ( Blattella germanica L.) avoids ingesting toxicant/diet mixtures because of an aversion to d -glucose in the diet rather than to the toxicant. d -fructose stimulated feeding in glucose-averse cockroaches. However, only fructose-glucose mixtures with a molar ratio ≥ 9:1 fructose:glucose stimulated feeding. Rather than being learned, glucose aversion is inherited as an autosomal incompletely dominant trait, which appears to be controlled by a single major gene. This is the first report of nutrient avoidance as a resistance mechanism and of aversion to this typically phagostimulatory molecule recognized as the universal metabolic fuel.

The diminutive supercolony: the Argentine ants of the southeastern United States

Native to Argentina and Brazil, the Argentine ant (Linepithema humile) is an invasive species that has become established on six continents and many oceanic islands. In several parts of its introduced range, including the western United States, southern Europe and Chile, the Argentine ant is unicolonial, forming extensive supercolonies. We examined population genetic structure and intercolony aggression in two regions of the introduced range of this species in the United States: California and the southeastern United States. Our results show that the southeastern L. humile population has high genotypic variability and strong intercolony aggression relative to the California population. In the California population, intercolony aggression was absent and 23 alleles were found across seven polymorphic microsatellite loci. However, in the Southeast, aggression between colonies was high and 47 alleles were present across the same seven loci in an equal number of colonies. We suggest that distinctly different colonization patterns for California and the Southeast may be responsible for the striking disparity in the genetic diversity of introduced populations. Southeastern colonies may have descended from multiple, independent introductions from the native range, undergoing a bottleneck at each introduction. In contrast, the California supercolony may have originated from one or more colonies inhabiting the southeastern United States, thus experiencing a double bottleneck. The differences in present‐day distribution patterns between California and the Southeast may be due to the combined effect of two factors: lower winter temperatures in the Southeast and/or competition with another successful and widely distributed ant invader, the fire ant Solenopsis invicta.

Diet-Related Modification of Cuticular Hydrocarbon Profiles of the Argentine Ant, Linepithema humile, Diminishes Intercolony Aggression

Territorial boundaries between conspecific social insect colonies are maintained through a highly developed nestmate recognition system modulated by heritable and, in some instances, nonheritable cues. Argentine ants, Linepithema humile, use both genetic and environmentally derived cues to discriminate nestmates from nonnestmates. We explored the possibility that intraspecific aggression in the Argentine ant might diminish when colonies shared a common diet. After segregating recently field-collected colony pairs into high or moderate aggression categories, we examined the effect of one of three diets: two hydrocarbon-rich insect prey, Blattella germanica and Supella longipalpa, and an artificial (insect-free) diet, on the magnitude of aggression loss. Aggression diminished between colony pairs that were initially moderately aggressive. However, initially highly aggressive colony pairs maintained high levels of injurious aggression throughout the study, independent of diet type. Each diet altered the cuticular hydrocarbon profile by contributing unique, diet-specific cues. We suggest that acquisition of common exogenous nestmate recognition cues from shared food sources may diminish aggression and promote fusion in neighboring colonies of the Argentine ant.

Acceptance and intake of gel and liquid sucrose compositions by the Argentine ant (Hymenoptera: Formicidae).

Abstract Liquids and gels are common delivery forms used in commercial ant baits, but the relative effectiveness of each is unknown. We compared the feeding responses of the Argentine ant, Linepithema humile (Mayr), to liquid and gel compositions of sucrose. In choice assays, more workers were counted on gel than liquid; however, substantially more liquid was consumed. Because workers could stand on the gel, more workers could feed simultaneously on the gel. The feeding bouts of individual workers, however, were much less efficient at extracting sucrose in gel form. Workers fed eightfold longer on the gel, yet removed fivefold less sucrose than workers feeding on liquid. This potential bias should be considered during attraction and palatability studies that use physically different bait compositions. When the toxicant fipronil was added to the compositions, a greater proportion of the colony died after workers had fed on liquid than gel baits. This finding suggests that liquid formulations may provide more effective control of Argentine ants due to the greater speed and abundance in which it is ingested.

Context-dependent nestmate discrimination and the effect of action thresholds on exogenous cue recognition in the Argentine ant

The optimal acceptance threshold model predicts that kin/nestmate discrimination is context dependent and that, in a fluctuating environment, the action component of nestmate discrimination is plastic, rather than static. We examined changes in intraspecific aggression among colonies of Argentine ants, Linepithema humile , in various discrimination contexts, and found that aggression occurred at higher rates when either nestmates or familiar territory indicated nest proximity, but not when social context was absent, thereby providing additional support for the optimal acceptance threshold model. Context-dependent aggression in the Argentine ant appears to result from a shift in acceptance threshold in response to fitness costs associated with accepting nonkin. The change in the action component of Argentine ant nestmate discrimination was explained to some degree by the hypothesis that the presence of nestmates indicates nest proximity and denotes a fitness payoff for active defence (nest indicator hypothesis) and by the hypothesis that nestmates share the cost of nest defence in groups, but not singly (cost minimizer hypothesis). Isolated nest referents (familiar territory, conspecific brood, or single familiar nestmates), however, had no effect on aggression thresholds. We provide mixed support for the hypothesis that workers from genetically less diverse colonies attack workers from more diverse colonies. We found that, in the context of nest defence, genetically more diverse colonies initiated attacks on colonies with lower genetic diversity. Therefore, the role of asymmetrical aggression in reducing genetic diversity within introduced populations of L. humile remains unknown and other extrinsic factors such as nest status and/or colony size may affect the outcome of aggressive interactions in the field. Finally, our finding that colonies reared under uniform conditions showed diminished intraspecific aggression only when assayed in a social and/or ecological context underscores the importance of using appropriate aggression assays for testing patterns of intercolony aggression in L. humile , and that the use of different rearing regimes, source colonies and collection times may produce contradictory results.

Hydrocarbon-released nestmate aggression in the Argentine ant, Linepithema humile, following encounters with insect prey

Argentine ants, Linepithema humile, were attacked by their nestmates following contact with a particular prey item, the brown-banded cockroach, Supella longipalpa. Contact with prey, as brief as 2 min, provoked nestmate aggression. Argentine ants contaminated with hydrocarbons extracted from S. longipalpa also released nestmate aggression behavior similar to that released by the whole prey item, confirming the involvement of hydrocarbons. In contrast to S. longipalpa, little or no nestmate aggression was induced by other ant prey from diverse taxa. A comparison of prey hydrocarbon profiles revealed that all hydrocarbons of S. longipalpa were very long chain components with 33 or more carbons, while other prey had either less, or none, of the very long chain hydrocarbons of 33 carbons or greater. We identified the hydrocarbons of S. longipalpa and some new groups of long chain hydrocarbons of L. humile. The majority of S. longipalpa hydrocarbons were 35 and 37 carbons in length with one to three methyl branches, and closely resembled two previously unidentified groups of compounds from L. humile of similar chain length. The hydrocarbons of S. longipalpa and L. humile were compared and their role in the Argentine ant nestmate recognition is discussed.

Urban areas may serve as habitat and corridors for dry-adapted, heat tolerant species; an example from ants

We collected ants from six urban and one forest land-use types in Raleigh, NC to examine the effects of urbanization on species richness and assemblage composition. Since urban areas are warmer (i.e., heat island effect) we also tested if cities were inhabited by species from warmer/drier environments. Species richness was lower in industrial areas relative to other urban and natural environments. There are two distinct ant assemblages; 1) areas with thick canopy cover, and 2) more disturbed open urban areas. Native ant assemblages in open environments have more southwestern (i.e., warmer/drier) distributions than forest assemblages. High native species richness suggests that urban environments may allow species to persist that are disappearing from natural habitat fragments. The subset of species adapted to warmer/drier environments indicates that urban areas may facilitate the movement of some species. This suggests that urban adapted ants may be particularly successful at tracking future climate change.

Geographical variation in Argentine ant aggression behaviour mediated by environmentally derived nestmate recognition cues

Social insects use a complex of recognition cues when discriminating nestmates from non-nestmate conspecifics. In the Argentine ant, Linepithema humile, recognition cues can be derived from exogenous sources, and L. humile acquires prey-derived hydrocarbons that are used in nestmate discrimination. We studied Argentine ant population-level distinctions in response to external recognition cues. Ants belonging to a California population were strongly affected by the imposition of prey-derived hydrocarbons, with spatially isolated colony fragments that had been fed different cockroach prey (Blattella germanica or Supella longipalpa) showing high and injurious intracolony aggression when reunited. In contrast, colonies of Argentine ants from the southeastern U.S. showed only moderate and noninjurious aggression when subjected to the same treatment. Field-collected colonies of L. humile had hydrocarbons in the range of those provided by S. longipalpa, and colonies from the southeastern U.S. had significantly higher initial levels of Supella-shared hydrocarbons. When fed cockroaches, Argentine ants from both regions acquired additional amounts of Supella- and Blattella-specific hydrocarbons, with a significant increase in levels of Blattella-specific hydrocarbons. Therefore, diet partitioning produced a greater change in the proportion of prey hydrocarbons in the California than in the southeastern U.S. populations, which may be responsible for the altered behaviour observed in the California population. Identifying factors underlying geographical variation in cue expression and/or perception may bring us closer to elucidating the selective forces driving nestmate recognition systems.