Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

Within-tree distribution of nest sites and foraging areas of ants on canopy trees in a tropical rainforest in Borneo

It has been argued that canopy trees in tropical rainforests harbor species-rich ant assemblages; however, how ants partition the space on trees has not been adequately elucidated. Therefore, we investigated within-tree distributions of nest sites and foraging areas of individual ant colonies on canopy trees in a tropical lowland rainforest in Southeast Asia. The species diversity and colony abundance of ants were both significantly greater in crowns than on trunks. The concentration of ant species and colonies in the tree crown seemed to be associated with greater variation in nest cavity type in the crown, compared to the trunk. For ants nesting on canopy trees, the numbers of colonies and species were both higher for ants foraging only during the daytime than for those foraging at night. Similarly, for ants foraging on canopy trees, both values were higher for ants foraging only during the daytime than for those foraging at night. For most ant colonies nesting on canopy trees, foraging areas were limited to nearby nests and within the same type of microhabitat (within-tree position). All ants foraging on canopy trees in the daytime nested on canopy trees, whereas some ants foraging on the canopy trees at night nested on the ground. These results suggest that spatial partitioning by ant assemblages on canopy trees in tropical rainforests is affected by microenvironmental heterogeneity generated by three-dimensional structures (e.g., trees, epiphytes, lianas, and aerial soils) in the crowns of canopy trees. Furthermore, ant diversity appears to be enriched by both temporal (diel) and fine-scale spatial partitioning of foraging activity.     

Mechanisms of dispersed central-place foraging in polydomous colonies of the Argentine ant

Many species of ants occupy multiple nests, a condition known as polydomy. Because of their decentralized structure, polydomous colonies may be removed from some of the constraints associated with classic central-place foraging. We used laboratory and field experiments to assess the mechanisms involved in dispersed central-place foraging in polydomous colonies of the Argentine ant Linepithema humile, a widespread invasive species. Both in the laboratory and in the field, Argentine ants established new nests at sites located near food. Laboratory colonies of L. humile redistributed workers, brood and resources among nests in response to the spatial heterogeneity of food resources. In addition, laboratory colonies formed recruitment trails between nests in the context of foraging, providing a mechanism for the transport of material between nests. This highly flexible system of allocating nests, workers and brood throughout a colony's foraging area potentially increases foraging efficiency and competitive ability. The importance of polydomy as a determinant of competitive ability is underscored by its prevalence among ecologically dominant ants, including most, if not all, highly invasive species. Copyright 2000 The Association for the Study of Animal Behaviour.     

Foraging preferences of ants on a heterogeneous Brazilian sandy shore habitat

1. Ants may select their food in response to nutritional needs of the colony and forage in a way that optimises a complementary nutrition. Even though resource availability is known to affect ant colony and individual health, there is still no study that has investigated the plastic preferences of ants according to spatial resource availability in naturally heterogeneous conditions. 2. Beaches are great biomes to test spatial foraging preference because a complete absence of nectaries can be found. Dorymyrmex nigra Pergande 1896 was found inhabiting a beach in southeastern Brazil, in which nectar sources are heterogeneously distributed. This study tested whether the foraging preference to sugar baits depended on the availability of nectar sources surrounding the nests. 3. We found that more D. nigra workers foraged on sugar baits when the colonies lacked naturally occurring nectar in their vicinity compared with colonies with abundant nectar nearby. 4. These results show that the foraging preference of ants depends upon resource availability. This is the first study to use a natural mosaic of resource availability to show that resource preference of ants is plastic and varies spatially.     

The Role of Non-Foraging Nests in Polydomous Wood Ant Colonies

A colony of red wood ants can inhabit more than one spatially separated nest, in a strategy called polydomy. Some nests within these polydomous colonies have no foraging trails to aphid colonies in the canopy. In this study we identify and investigate the possible roles of non-foraging nests in polydomous colonies of the wood ant Formica lugubris. To investigate the role of non-foraging nests we: (i) monitored colonies for three years; (ii) observed the resources being transported between non-foraging nests and the rest of the colony; (iii) measured the amount of extra-nest activity around non-foraging and foraging nests. We used these datasets to investigate the extent to which non-foraging nests within polydomous colonies are acting as: part of the colony expansion process; hunting and scavenging specialists; brood-development specialists; seasonal foragers; or a selfish strategy exploiting the foraging effort of the rest of the colony. We found that, rather than having a specialised role, non-foraging nests are part of the process of colony expansion. Polydomous colonies expand by founding new nests in the area surrounding the existing nests. Nests founded near food begin foraging and become part of the colony; other nests are not founded near food sources and do not initially forage. Some of these non-foraging nests eventually begin foraging; others do not and are abandoned. This is a method of colony growth not available to colonies inhabiting a single nest, and may be an important advantage of the polydomous nesting strategy, allowing the colony to expand into profitable areas.     

Soil surface complexity has a larger effect on food exploitation by ants than a change from grassland to shrubland

1. Changes in vegetation community composition, such as a transition from grassland to shrubland (woody encroachment), are associated with reductions in plant cover and increases in bare ground. Encroachment‐driven changes in surface cover at small spatial scales can alter ant community assemblages by changing their foraging behaviour and their ability to locate and monopolise resources. 2. Artificial arenas with three levels of complexity were used to examine changes in ant foraging efficiency, body size and ability to monopolise food. The three levels of complexity included a control (no substrate), low‐complexity treatment (woody debris) and high‐complexity treatment (leaf litter). 3. No difference was found in ant species composition within the complexity arenas between grassland and shrubland, but ant functional groups ‘generalised Myrmicinae’ and ‘subordinate Camponotini’ were more abundant in grassland arenas, whereas ‘opportunists’ were more abundant in shrubland arenas. Ants took twice as long to find baits in high‐complexity treatments, and 1.5 times as long in low‐complexity treatments, than in control treatments, which were bare arenas with no substrate. Ant body size declined with increasing surface complexity, suggesting that larger ants are discouraged from foraging in complex habitats. 4. There was also significantly greater monopolisation of the protein bait (tuna) in low‐ and high‐complexity treatments, but there were no differences between tuna and carbohydrate (honey) in the control treatment. Consistently, no differences were found in ant behaviour between grasslands and shrublands. 5. The present study shows that ants are more responsive to small‐scale alterations in soil surface complexity than to changes in vegetation community composition. Changes in soil surface complexity select for ants based on body size, which in turn influences their foraging success. Changes in vegetation complexity at small spatial scales are therefore likely to influence ant behaviour and abundance of some functional groups, potentially having an effect on the many ecosystem functions carried out by ants.     

The size–distance relationship in the wood ant Formica rufa

1. The size–distance relationship among honeydew‐collecting foragers of the red wood ant Formica rufa was investigated. Within the colony territory, the size (as measured by head width) and fresh weight of samples of foragers were determined for ants ascending and descending trees near, and farther from, the central nest mound. 2. The mean size of the ants was significantly higher at far trees than at near trees in six out of the seven colonies investigated, confirming the general presence of the size–distance relationship. 3. In three colonies, a load–distance relationship was also found. For a given head width, honeydew‐carrying ants descending far trees were significantly heavier than those descending near trees (i.e. they were carrying heavier loads from trees farther away from the central nest mound). 4. This is the first time that both load–distance and size–distance relationships have been reported in foraging workers from the same ant colony. 5. The combined effects of these characteristics suggest that colony foraging efficiency is enhanced by far trees being visited by the larger workers that then return with heavier loads of honeydew.     

Nest connectivity and colony structure in unicolonial Argentine ants

Unicolonial ant colonies occupy many nests and individuals rarely show aggression across large geographic distances. These traits make it difficult to detect colony structure. Here we identify colony structure at scales of hundreds of square-meters, within an invasive population of unicolonial Argentine ants. In experiments using labeled food, and in a 3-year census of nests and trails, we found that food was shared and nests were linked by trails at distances up to 50 meters. Food was not distributed to all nearby Argentine ant nests, showing that ants tend to share resources within a spatially bounded group of nests. The spatial extent of food sharing increased from winter to summer. Across different habitats and nest densities, nests were consistently aggregated at spatial scales of 3- 4 meters in radius. This suggests that new nests bud from old nests at short distances regardless of local conditions. We suggest that a ‘colony’ of Argentine ants could be defined as a group of nests among which ants travel and share food. In our study population, colonies occupy up to 650 m² and contain as many as 5 million ants. In combination with previous work showing that there is genetic differentiation among nests at similar spatial scales, the results suggest that Argentine ant populations do not function ecologically as single, large supercolonies, but instead as mosaics of smaller, distinct colonies consisting of groups of interacting nests. Received 6 June 2008; revised 30 June 2008; accepted 2 July 2008.     

Carbohydrate as Fuel for Foraging,Resource Defense and Colony Growth – a Long‐term Experiment with the Plant‐ant Crematogaster nigriceps

Mismatches in nutrient composition (e.g., protein, carbohydrates, lipids, etc.) between consumers and the resources they depend on can have ecological consequences, affecting traits from individual behavior to community structure. In many terrestrial ecosystems, ants depend on plant and insect mutualist partners for carbohydrate‐rich rewards that are nutritionally unbalanced (especially in protein) relative to colony needs. Despite imbalances, many carbohydrate‐feeding ant mutualists dominate communities—both competitively and numerically—raising the question of whether excess carbohydrates ‘fuel’ colony acquisition of limiting resources and growth. In a 10‐month field study, we manipulated carbohydrate access for the obligate plant‐ant Crematogaster nigriceps to test whether carbohydrate availability could be mechanistically linked to ecological dominance via heightened territory defense, increased protein foraging, and colony growth. Supplementation increased aggressive defense of hosts after only two weeks, but was also strongly linked to variation in rainfall. Contrary to predictions, we did not find that supplemented colonies increased protein foraging. Instead, colonies with reduced carbohydrate access discovered a greater proportion of protein baits, suggesting that carbohydrate deprivation increases foraging intensity. We found no significant effect of carbohydrate manipulation on brood or alate production. These results contrast with findings from several recent short‐term and lab‐based nutrient supplementation studies and highlight the role of seasonality and biotic context in colony‐foraging and reproductive decisions. These factors may be essential to understanding the consequences of carbohydrate access in natural plant‐ant systems.     

Colony structure and spatial distribution of food resources in the polydomous meat ant <Emphasis Type="Italic">Iridomyrmex purpureus</Emphasis>

Polydomous social insects may reduce the costs of foraging by the strategic distribution of nests throughout their territory or home-range. This efficiency may most likely be achieved if the resources are relatively stable in place and time, and the colonies and nests are distributed in response to the location of the resources. However, no study has investigated how the distribution of food sources influences the spatial patterns of nests within polydomous colonies under natural conditions. Our two year study of 140 colonies of the Australian ant Iridomyrmex purpureus revealed that the decentralization of nests within colonies is associated with the distribution of trees containing honey-dew producing hemiptera. We show there is a positive correlation between the maximum distance between trees containing hemiptera and the maximum distance between nests within a colony. In addition, we demonstrate the mechanism by which this pattern may arise; new nests are built nearer to trees containing hemiptera than existing nests. Further, the distance between trees containing hemiptera and the nearest nests was negatively correlated with the length of exploitation of that tree. Finally, we show that most food is delivered to the nearest nest after which other ants redistribute it between the nests. Combined, these data suggest that foraging efficiency may be an important selection pressure favouring polydomy in I. purpureus. Received 6 April 2006; revised 29 September; accepted 4 October 2006.     

Northward expansion of the invasive Linepithema humile (Hymenoptera: Formicidae) in the eastern United States is constrained by winter soil temperatures

The invasive Argentine ant, Linepithema humile (Mayr) (Hymenoptera: Formicidae) has been evident in the North Carolina Piedmont, United States for 90 yr but has failed to spread further north. We investigated the mechanisms preventing this expansion. The Argentine ant ceases foraging at temperatures below 5°C and we hypothesized that winter soil temperatures at higher latitudes restricted foraging long enough to cause colony starvation. We tested if the Argentine ant could successfully feed at temperatures below 5°C and found that colonies would starve. We subjected Argentine ant nests to a range of sub- and above-freezing temperatures and measured worker mortality at various time intervals. We found that Argentine ant colonies will collapse after 8.5 d at 5°C. Argentine ants can escape ambient cold temperatures by moving nests into the soil column. We tested how deeply into the soil Argentine ant queens and workers need to move to survive winter in North Carolina. Soil temperatures in the North Carolina Piedmont do not fall below 5°C for longer than nine consecutive days; therefore, Argentine ant colonies need only to retreat a few centimeters into the soil column to escape unsuitable temperatures. Winter soil temperature data from four climate stations situated from latitudes 35°, the current Eastern United States latitudinal limit for Argentine ant population expansion, to 39° were searched for periods where soil temperatures would have led to colony extirpation. North of their current distributions, extended periods of soil temperatures below 5°C regularly occur, preventing Argentine ant colonies from persisting.     

Strong influences of a dominant,ground‐nesting ant on recruitment,and establishment of ant colonies and communities

Many factors drive the organization of communities including environmental factors, dispersal abilities, and competition. In particular, ant communities have high levels of interspecific competition and dominance that may affect community assembly processes. We used a combination of surveys and nest supplementation experiments to examine effects of a dominant ground‐nesting ant (Pheidole synanthropica) on (1) arboreal twig‐nesting, (2) ground‐foraging, and (3) coffee‐foraging ant communities in coffee agroecosystems. We surveyed these communities in high‐ and low‐density areas of P. synanthropica over 2 years. To test for effects on twig ant recruitment, we placed artificial nesting resources on coffee plants in areas with and without P. synanthropica. The first sampling period revealed differences in ant species composition on the ground, in coffee plants, and artificial nests between high‐ and low‐density sites of P. synanthropica. High‐density sites also had significantly lower recruitment of twig ants and had species‐specific effects on twig ant species. Prior to the second survey period, abundance of P. synanthropica declined in the high‐density sites, such that P. synanthropica densities no longer differed. Subsequent sampling revealed no difference in total recruitment of twig ants to artificial nests between treatments. Likewise, surveys of ground and coffee ants no longer showed significant differences in community composition. The results from the first experimental period, followed by survey results after the decline in P. synanthropica densities suggest that dominant ants can drive community assembly via both recruitment and establishment of colonies within the community.     

The Regulation of Ant Colony Foraging Activity without Spatial Information

Many dynamical networks, such as the ones that produce the collective behavior of social insects, operate without any central control, instead arising from local interactions among individuals. A well-studied example is the formation of recruitment trails in ant colonies, but many ant species do not use pheromone trails. We present a model of the regulation of foraging by harvester ant (Pogonomyrmex barbatus) colonies. This species forages for scattered seeds that one ant can retrieve on its own, so there is no need for spatial information such as pheromone trails that lead ants to specific locations. Previous work shows that colony foraging activity, the rate at which ants go out to search individually for seeds, is regulated in response to current food availability throughout the colony's foraging area. Ants use the rate of brief antennal contacts inside the nest between foragers returning with food and outgoing foragers available to leave the nest on the next foraging trip. Here we present a feedback-based algorithm that captures the main features of data from field experiments in which the rate of returning foragers was manipulated. The algorithm draws on our finding that the distribution of intervals between successive ants returning to the nest is a Poisson process. We fitted the parameter that estimates the effect of each returning forager on the rate at which outgoing foragers leave the nest. We found that correlations between observed rates of returning foragers and simulated rates of outgoing foragers, using our model, were similar to those in the data. Our simple stochastic model shows how the regulation of ant colony foraging can operate without spatial information, describing a process at the level of individual ants that predicts the overall foraging activity of the colony.     

Behavioral mechanisms underlying ants' density-dependent deterrence of aphid-eating predators

Density-dependent mutualisms have been well documented, but the behavioral mechanisms that can produce such interactions are not as well understood. We investigated interactions between predatory ladybirds and the ant Lasius neoniger, which engages in a facultative association with the aphid Aphis fabae . We found that ants disrupted predator aggregation and deterred foraging, but that this effect varied with aphid density. In the field, smaller aphid colonies had higher numbers of ants per aphid (higher relative ant density), whereas plants with larger aphid colonies had lower relative ant density. Ants deterred ladybird foraging when relative ant density was high, but when relative ant density was low, ladybirds aggregated to aphids and foraged more successfully. This difference in ladybird foraging success appeared to be driven by variation in the ants' distribution on the plant and the ladybirds' reaction to ants. When relative ant density was high, ants moved around the perimeter of the aphid colonies, which resulted in faster detection of predators and a greater likelihood of ladybirds leaving. However, when relative ant density was low, ants moved only in the midst of the aphid colonies and rarely around the perimeter, which allowed predators to approach the aphid colony from the perimeter and feed without detection. Such predators were less likely to leave the aphid colony when subsequently detected by ants. We suggest that differences in relative ant numbers, ant distribution, and predator reaction to detection by ants could lead to complex population-level consequences including density-dependent mutualisms and the possibility that predators act as prudent predators.     

Intraspecific Food‐Robbing and Neighborhood Competition: Consequences for Anti‐Robber Vigilance and Colony Productivity

Most social animals have mechanisms to distinguish group members from outsiders, in part to prevent the exploitation of resources reserved for members of the group. Nevertheless, specialized thieves of the Neotropical ant, Ectatomma ruidum, also known as the ‘thieving ant’, regularly enter and steal resources from distinct, neighboring colonies. Here, we examine the mechanisms and consequences of thievery in a population of E. ruidum. We show that (1) individuals from nearby colonies were accepted more often than those from farther colonies; (2) rejection rates decreased as individuals interacted more with non‐nestmates from the same source colony; and (3) colonies that were experimentally treated to reduce thievery rates had improved productivity. The boost in productivity with thievery reduction was greater in low density populations than in high density populations. We conclude that, as in other species, thievery has negative fitness costs to E. ruidum. However, greater acceptance of neighbors than non‐neighbors and increased acceptance after habituation to non‐nestmates suggest a proximate explanation for the presence of thievery. Moreover, lower fitness costs of thievery at high nesting density, combined with observations of extraordinarily high densities of E. ruidum throughout its range, suggest there is little selection pressure among these ants to guard against thieves, thus providing an ultimate explanation why thievery persists among litter‐foraging ants.     

Specificity and stability of the Acromyrmex–Pseudonocardia symbiosis

The stability of mutualistic interactions is likely to be affected by the genetic diversity of symbionts that compete for the same functional niche. Fungus‐growing (attine) ants have multiple complex symbioses and thus provide ample opportunities to address questions of symbiont specificity and diversity. Among the partners are Actinobacteria of the genus Pseudonocardia that are maintained on the ant cuticle to produce antibiotics, primarily against a fungal parasite of the mutualistic gardens. The symbiosis has been assumed to be a hallmark of evolutionary stability, but this notion has been challenged by culturing and sequencing data indicating an unpredictably high diversity. We used 454 pyrosequencing of 16S rRNA to estimate the diversity of the cuticular bacterial community of the leaf‐cutting ant Acromyrmex echinatior and other fungus‐growing ants from Gamboa, Panama. Both field and laboratory samples of the same colonies were collected, the latter after colonies had been kept under laboratory conditions for up to 10 years. We show that bacterial communities are highly colony‐specific and stable over time. The majority of colonies (25/26) had a single dominant Pseudonocardia strain, and only two strains were found in the Gamboa population across 17 years, confirming an earlier study. The microbial community on newly hatched ants consisted almost exclusively of a single strain of Pseudonocardia while other Actinobacteria were identified on older, foraging ants in varying but usually much lower abundances. These findings are consistent with recent theory predicting that mixtures of antibiotic‐producing bacteria can remain mutualistic when dominated by a single vertically transmitted and resource‐demanding strain.     

Field study on the foraging characteristics of a ponerine ant,Hagensia havilandi Forel

Summary A field study of the foraging strategy used by the ponerine ant,Hagensia havilandi is reported. They have permanent nests in the leaf litter of coastal forests.H. havilandi is a diurnal forager and collects a variety of live and dead arthropods. These predatory ants exhibit individual foraging with no cooperation in the search for or retrieval of food items. Three colonies were observed and showed similar temporal and spatial foraging patterns. The paths of individual ants were followed and the results showed that the foragers exhibit area fidelity, and return to the nest via a direct route on finding on prey item. Several foragers did not return to the nest at dusk but returned the following morning. Occasionally a limited amount of tandem recruitment was displayed.     

Parallel foraging cycles for different resources in leaf‐cutting ants: a clue to the mechanisms of rhythmic activity

1. Leaf‐cutting ants display regular diel cycles of foraging, but the regulatory mechanisms underlying these cycles are not well known. There are, however, some indications in the literature that accumulation of leaf tissue inside a nest dampens recruitment of foragers, thereby providing a negative feedback that can lead to periodic foraging. We investigated two foraging cycles occurring simultaneously in an Atta colombica colony, one involving leaf harvesting and the other exploiting an ephemeral crop of ripe fruit. 2. Leaf harvesting followed a typical diel pattern of a 10–12 h foraging bout followed by a period of inactivity, while fruit harvesting occurred continuously, but with a regular pre‐dawn dip in activity that marked a 24 h cycle. 3. Although the results of the present study are drawn from a single field colony, the difference found is consistent with a mechanism of negative feedback regulation acting in parallel on two resources that differ in their rates of distribution and processing, creating cycles of formation and depletion of material caches. 4. This hypothesis should provoke further interest from students of ant behaviour and some simple manipulative experiments that would begin to test it are outlined. Any role of resource caches in regulating foraging by Atta colonies may have similarities to the logistics of warehouse inventories in human economic activity.     

Effect of harvester ants of the genus Pogonomyrmex on the soil seed bank around their nests in the central Monte desert,Argentina

1. The abundance and composition of soil seed banks is a key determinant of plant community structure. Harvester ants can remove huge quantities of preferred seeds close to the nest affecting composition and spatial distribution of plants. 2. In the central Monte desert (Argentina) ants of the genus Pogonomyrmex have high seed removal rates, especially of the five main grasses. The aim of this study was to establish if their foraging activity affects spatial patterns of the soil seed bank around their nests. Our hypotheses were: (1) removal by ants decreases seed abundance of preferred species in the soil; and (2) the effect varies in time. 3. Soil seed abundance was assessed at different distances from Pogonomyrmex nests in the litter and in bare soil at the beginning, the middle, and the end of the season (late spring‐early autumn). 4. A lower seed abundance of preferred species was observed close to the nest in the litter at the end of the season. Non‐preferred species showed no distance gradient. 5. The lower foraging activity and seed consumption at the beginning of the season could explain the temporal variation of the spatial effect. This was only observed in the litter, probably because of the higher removal frequency in this substrate. 6. Colonies of Pogonomyrmex spp. could enhance the heterogeneity of soil seed banks in the central Monte desert from the summer to the beginning of the autumn. Implications for vegetation dynamics depend on the degree to which seed density limits perennial grasses recruitment after ant activity season.     

Litter Biomass and Nutrient Determinants of Ant Density, Nest Size, and Growth in a Costa Rican Tropical Wet Forest

The nutritional demands of animals vary by taxon. Across landscapes, communities of animals experience variability in the stoichiometry of carbon and nutrients within their resource base. Thus, we expect stoichiometry to contribute to the spatial variance in the demographic parameters of animal communities. Here, we measure how the composition of a litter-nesting tropical rainforest ant community is influenced by spatial variation in environmental stoichiometry relative to litter biomass, a known predictor of ant density. We found the density of ants and their nests were strongly related to litter biomass and carbon: phosphorus stoichiometry. The spatial variation in soil nutrients, which determines leaf litter stoichiometry, was an excellent predictor of nest size in the two most common genera of ants. We found a negative relationship between species' growth rate and local soil stocks of phosphorus. Overall, the density of litter-dwelling ants varied greatly across this tropical forest landscape and environmental stoichiometry can account for limits on ant density independent of the biomass of the leaf litter resource base.