Nest raiding by the invasive Argentine ant on colonies of the harvester ant, Pogonomyrmex subnitidus

Invasive ants often displace native ants, and published studies that focus on these interactions usually emphasize interspecific competition for food resources as a key mechanism responsible for the demise of native ants. Although less well documented, nest raiding by invasive ants may also contribute to the extirpation of native ants. In coastal southern California, for example, invasive Argentine ants (Linepithema humile) commonly raid colonies of the harvester ant, Pogonomyrmex subnitidus. On a seasonal basis the frequency and intensity of raids vary, but raids occur only when abiotic conditions are suitable for both species. In the short term these organized attacks cause harvester ants to cease foraging and to plug their nest entrances. In unstaged, one-on-one interactions between P. subnitidus and L. humile workers, Argentine ants behaved aggressively in over two thirds of all pair-wise interactions, despite the much larger size of P. subnitidus. The short-term introduction of experimental Argentine ant colonies outside of P. subnitidus nest entrances stimulated behaviors similar to those observed in raids: P. subnitidus decreased its foraging activity and increased the number of nest entrance workers (many of which labored to plug their nest entrances). Raids are not likely to be the result of competition for food. As expected, P. subnitidus foraged primarily on plant material (85% of food items obtained from returning foragers), but also collected some dead insects (7% of food items). In buffet-style choice tests in which we offered Argentine ants food items obtained from P. subnitidus, L. humile only showed interest in dead insects. In other feeding trials L. humile consistently moved harvester ant brood into their nests (where they were presumably consumed) but showed little interest in freshly dead workers. The raiding behavior described here obscures the distinction between interspecific competition and predation, and may well play an important role in the displacement of native ants, especially those that are ecologically dissimilar to L. humile with respect to diet. Received 15 July 2005; revised 19 October 2005; accepted 26 October 2005.     

The effect of neighbours on the mortality of harvester ant colonies

1. This study investigates how the density and size of conspecific neighbours affected the mortality of established colonies. Data were from a population of 250–300 colonies of the red harvester ant [ Pogonomyrmex barbatus (F. Smith)], on a 10-ha site, from 1988 to 1995.
2. Colonies are more likely to die as they grow older.
3. Small neighbours tend to lower the probability of dying.
4. The probability of dying varied significantly from year to year.
5. There was little effect of crowding on mortality.
6. There appeared to be no spatial effects, on the scale of 25. m, on the probability of dying.
7. Previous work shows effects of density on the founding of new colonies and on the reproductive output of established ones. The results presented here indicate that competition with neighbours rarely causes the death of established colonies.     

Nest relocation and encounters between colonies of the seed-harvesting ant Messor andrei

Summary: Nest relocation in ants may be an attempt to escape areas of high competition. Encounters between colonies have been suggested to be the proximate cause of nest relocation. I examined the relation between nest relocation and encounters in a population of the seed-harvesting ant Messor andrei, a native of the western United States. Over 80% of colonies relocated their nest site during a year. Colonies moved up to ten times in a year, and new nest-sites were more distant from their nearest neighbour. However, encounters did not precipitate nest relocation. Relocation was not more likely to occur directly after an encounter, and colonies that moved did not experience more encounters than other colonies. Other possible cues for nest relocation, including predation, disease, microclimatic effects, and local resource depletion, are discussed.     

The regulation of foraging activity in red harvester ant colonies

Behavioral plasticity in social insects is intriguing because colonies adjust to environmental change through the aggregated responses of individuals. Without central control, colonies adjust numbers of workers allocated to various tasks. Individual decisions are based on local information from the environment and other workers. This study examines how colonies of the seed-eating ant Pogonomyrmex barbatus adjust the intensity of foraging in an arid environment where conspecific neighbors compete for foraging area. The main question is how foragers decide whether to leave the nest. Patrollers search the area before foragers emerge. Removal experiments show that the return of the patrollers stimulates the onset of foraging, and later, the rate at which foragers return affects the rate at which foragers continue to leave the nest. Foraging activity is less sensitive to changes in the rate of returning foragers than to changes in the rate of returning patrollers. These results suggest that whether a colony forages at all on a given day depends on conditions detected early by patrollers but that once foraging begins, the intensity of foraging does not track, on an hourly timescale, how quickly foragers can find food.     

Nest site selection by fire ant queens

Summary We examined nest site selection by foundresses of the polygyne form ofSolenopsis invicta. In the laboratory, foundresses were allowed to nest in control soil vs. soil inoculated with refuse from a colony ofPheidole dentata, a natural enemy. In a second experiment foundresses chose between control andS. invicta soil. More foundresses (100%) avoided the soil of a natural enemy than the soil of a conspecific colony (79%) that may accept them into the nest. Foundresses from this polygyne ant appear to avoid other colonies in general, but avoid heterospecific colonies more.     

Nest architecture in the western harvester ant,Pogonomyrmex occidentalis (Cresson)

Summary The structure and thermal environment produced by the nest cones of the western harvester ant,Pogonomyrmex occidentalis, are investigated. The nest cone is oriented so that the longest slope faces towards the southeast ad the nest entrance faces the southeast. The temperature of the soil was monitored at thirty-six locations within the nest in order to measure the daily temperature change as a function of depth, aspect and radial distance from the center of peak of the cone. The occupancy of the nest cone by workers and by brood was assessed by core samples taken at different times of day in different portions of the cone. The brood are only present during the midmorning sample and are present in greatest numbers on the eastern side of the cone. The adult workers are present in the cone in highest numbers during the midmorning, virtually absent from the cone during mid-afternoon and returning in smaller numbers at dusk, apparently spending the night near the surface on the eastern side of the mound. The eastern and southern portions of the cone are occupied most frequently while the western side of the cone is nearly vacant. The nest cone is an adaptation which magnifies the amplitude of daily temperature fluctation. By moving to appropriate areas of the nest cone, the ants can take advantage of the increased range of temperature.     

Nest site limitation and colony takeover in the ant Leptothorax nylanderi

Ecological constraints on the success of independent coloniesare thought to strongly shape the organization of ant societies.One of the most important factors is probably the availabilityof suitable empty nest sites. By population censuses, laboratoryexperiments, and microsatellite analyses, we investigated thecolony and population structure of the small, myrmicine antLeptothorax (Myrafant) nylanderi in a deciduous forest nearWürzburg, Germany, where nest sites appear to be stronglylimited, especially in late summer. Colonies of L. nylanderiinhabit cavities in rotting branches, hollow acorns, grass stems,etc. After hibernation, a temporary overabundance of empty nestsites facilitates the fragmentation of larger colonies intosmaller buds, which, because the species is monogynous, arein part queenless. Nest sites become scarce in summer due torapid deca and both established colonies and young foundingqueens face a severe shortage of suitable nest sites. This leadsto the fusion of established, unrelated colonies, which afterinitial fighting permanently merge and live together. Typicalyonly one queen survives after fusion. Similarly, young matedqueens may seek adoption in alien nests instead of foundingtheir own colonies solitarily, and here again only a singlequeen survives. This temporary intraspecific parasitism maybe an important first step in the evolution of obligatory permanentparasitism, which is widespread in the genus Leptothorax.     

The relative availabilities of complementary resources affect the feeding preferences of ant colonies

Theory predicts that consumers selecting among complementaryresources will show stronger preferences for items that becomerelatively less available. I tested this hypothesis in a fieldstudy that compared the preferences of ant colonies given simultaneousaccess to experimental foods differing in carbohydrate and proteincontent. In the first part of the study, I examined the effectof nutrient supplementation on colony-level preference in theant Dorymyrmex smithi. Colonies that had received a proteinsolution for 24 h consumed proportionally more carbohydratesthan control colonies that had been given access to water, suggestingthat colonies preferred nutrients when they became relativelyrare. In the second part of the study, I compared colony-levelpreference among eight species of ants that differ in theirrelative access to carbohydrates and protein in the field. Ifound that species with relatively easy access to carbohydratespreferred protein, whereas species with greater access to proteinpreferred carbohydrates. These results suggest that the benefitsof a nutritionally mixed diet coupled with differences in therelative availability of nutrients may explain variation infeeding decisions both within and among ant species.     

Foraging arena size and structural complexity affect the dynamics of food distribution in ant colonies

Food acquisition by ant colonies is a complex process that starts with acquiring food at the source (i.e., foraging) and culminates with food exchange in or around the nest (i.e., feeding). While ant foraging behavior is relatively well understood, the process of food distribution has received little attention, largely because of the lack of methodology that allows for accurate monitoring of food flow. In this study, we used the odorous house ant, Tapinoma sessile (Say) to investigate the effect of foraging arena size and structural complexity on the rate and the extent of spread of liquid carbohydrate food (sucrose solution) throughout a colony. To track the movement of food, we used protein marking and double-antibody sandwich enzyme-linked immunosorbent assay, DAS-ELISA. Variation in arena size, in conjunction with different colony sizes, allowed us to test the effect of different worker densities on food distribution. Results demonstrate that both arena size and colony size have a significant effect on the spread of the food and the number of workers receiving food decreased as arena size and colony size increased. When colony size was kept constant and arena size increased, the percentage of workers testing positive for the marker decreased, most likely because of fewer trophallactic interactions resulting from lower worker density. When arena size was kept constant and colony size increased, the percentage of workers testing positive decreased. Nonrandom (clustered) worker dispersion and a limited supply of food may have contributed to this result. Overall, results suggest that food distribution is more complete is smaller colonies regardless of the size of the foraging arena and that colony size, rather than worker density, is the primary factor affecting food distribution. The structural complexity of foraging arenas ranged from simple, two-dimensional space (empty arenas) to complex, three-dimensional space (arenas filled with mulch). The structural complexity of foraging arenas had a significant effect on food distribution and the presence of substrate significantly inhibited the spread of food. Structural complexity of foraging arenas and the resulting worker activity patterns might exert considerable influence on socioecological processes in ants and should be considered in laboratory assays.     

Characterizing the collective personality of ant societies: aggressive colonies do not abandon their home

Animal groups can show consistent behaviors or personalities just like solitary animals. We studied the collective behavior of Temnothorax nylanderi ant colonies, including consistency in behavior and correlations between different behavioral traits. We focused on four collective behaviors (aggression against intruders, nest relocation, removal of infected corpses and nest reconstruction) and also tested for links to the immune defense level of a colony and a fitness component (per-capita productivity). Behaviors leading to an increased exposure of ants to micro-parasites were expected to be positively associated with immune defense measures and indeed colonies that often relocated to other nest sites showed increased immune defense levels. Besides, colonies that responded with low aggression to intruders or failed to remove infected corpses, showed a higher likelihood to move to a new nest site. This resembles the trade-off between aggression and relocation often observed in solitary animals. Finally, one of the behaviors, nest reconstruction, was positively linked to per-capita productivity, whereas other colony-level behaviors, such as aggression against intruders, showed no association, albeit all behaviors were expected to be important for fitness under field conditions. In summary, our study shows that ant societies exhibit complex personalities that can be associated to the physiology and fitness of the colony. Some of these behaviors are linked in suites of correlated behaviors, similar to personalities of solitary animals.     

Nest site selection by the Argentine ant and suitability of artificial nests as a control tool

The Argentine ant is an invasive species that has been introduced worldwide causing devastating effects on entire ecosystems. Control strategies might be focused on slowing its rate of spread to limit its establishment inside yet non-invaded areas. For this, a better knowledge about nest selection is necessary to identify rapidly and accurately nest locations where to apply control measures. Herein, nest site selection by the Argentine ant, nests’ physical characteristics and their longevity were studied in three invaded cork oak secondary forest. Results showed that this species shifts nest locations seasonally to keep appropriate microclimatic conditions, nesting mainly underneath rocks during cold and rainy months and in tree bases during warmer periods. The terrain features at micro-scale (orientation and slope) were found to influence the distribution of the Argentine ant nests beneath rocks. Additionally, artificial nests used as a control tool were tested, finding that their use may be suitable if they are set in appropriate locations and before the ants start migrating to winter aggregations.     

Nest architecture shapes the collective behaviour of harvester ants

Structures influence how individuals interact and, therefore, shape the collective behaviours that emerge from these interactions. Here I show that the structure of a nest influences the collective behaviour of harvester ant colonies. Using network analysis, I quantify nest architecture and find that as chamber connectivity and redundancy of connections among chambers increase, so does a colony''s speed of recruitment to food. Interestingly, the volume of the chambers did not influence speed of recruitment, suggesting that the spatial organization of a nest has a greater impact on collective behaviour than the number of workers it can hold. Thus, by changing spatial constraints on social interactions organisms can modify their behaviour and impact their fitness.     

Nest site selection and longevity in the ponerine ant Rhytidoponera metallica (Hymenoptera, Formicidae)

Summary: The size of the nesting cavity and the nature of the nest-building material can constrain colony growth in eusocial insects. Rocks protect colony members against extreme temperatures as well as serve as a supplementary source of heat, thereby maintaining optimal conditions for brood development. The frequency distribution of available rocks and those occupied by colonies of Rhytidoponera metallica in the Avon River State Forest, Victoria, Australia, showed that colonies of R. metallica avoided nesting under small rocks (<200 cm²). A strong correlation between colony size and rock size indicates that nesting under a larger rock promotes colony growth. Laboratory choice experiments revealed that workers are capable of recognizing larger rocks from external physical characters (dimension). Field data indicates that colonies leave nests frequently, with nest abandonment peaking in the summer months.     

Nest relocation in the ant Pheidole dentata

We studied nest relocation in the ant Pheidole dentata, a common species in the southern US, by following colonies for 6 weeks. We correlated probability of relocation with several abiotic and biotic environmental factors, such as air temperature, humidity, leaf litter depth (LLD), nest type and presence of phorid fly parasitoids. Colonies moved often, on average every 16.2 days. By the end of our study, fewer than 5 % of colonies marked at the start remained in their original nests. Only 3.1 % of colonies returned to a previously used nest. The mean distance of relocation was 0.76 ± 0.73 m. Colonies nested in a variety of locations, such as in the ground, by tree trunks, under rocks and inside wood. Several factors affected probability of relocation. Higher LLD and vapor pressure deficit (VPD) increased probability of relocation. Probability of relocation varied by nest type, by plot and week of study. We found no evidence that surface cover or rain affected relocation behavior. Colonies appear to avoid nests that become dry by nesting against tree trunks. Phorid fly abundance correlated negatively with VPD, yet it had no effect on colonies’ likelihood of relocation. We discuss other implications of frequent nest relocations.     

Nest defense behavior against ant attacks in post-emergent colonies of wasp Mischocyttarus cerberus (Hymenoptera, Vespidae)

The primitive eusocial wasp Mischocyttarus cerberus forms colonies of independent foundation, without morphological differentiation among castes. Ants are natural enemies of the social wasps and defending the wasps’ nests involves chemical and active defense strategies. The aims of this work were to verify the kind of defense the wasps use most frequently in post-emergent colonies of M. cerberus. We also observed whether the nest was abandoned during the ant attack and whether any relationship existed between the forms that colony defense took and the number of adults, the number of cells, and the number of immature wasps. The study was carried out on the campus of Universidade Estadual Paulista of Rio Claro, São Paulo, Brazil. The 23 nests under study were mapped weekly, and 68 bioassays were performed by simulating ant attacks against the nests. The results showed that wasps used both active and chemical strategies for nest defense, and the PCA analysis showed that the aggressive behaviors of biting the ant, wings vibrating, gaster hitting, and abdomen pumping were the dominant terms; the PCA correlation values were 2.70, 2.54, 1.64, and 1.63, respectively. The colonies in pre- and post-male substages with few immature wasps and the nests in post-male substage with one adult were more correlated with the nonaggressive behaviors of hiding, staying immobile, and flying; their PCA correlation values were 3.12, 2.56, and 1.77, respectively. These results show that the number of immature wasps is an important factor in the kind and in the intensity of the defense behavior against ant attacks.     

Recruitment pheromone in the harvester ant genus Pogonomyrmex

Workers of the harvester ant genus Pogonomyrmex employ recruitment trail pheromones discharged from the poison gland. In P. barbatus, P. maricopa, P. occidentalis and P. rugosus we identified three pyrazines [2,5-dimethylpyrazine, trimethylpyrazine and 3-ethyl-2,5-dimethylpyrazine (EDMP)] as major compounds of the volatile part of the poison-gland secretions. Laboratory and field tests revealed EDMP to be the main recruitment pheromone.