Macronutrient regulation in the Rasberry crazy ant (<Emphasis Type="Italic">Nylanderia</Emphasis> sp. nr. <Emphasis Type="Italic">pubens</Emphasis>)

Animals grow and optimize performance when they collect foods in amounts and ratios that best meet their species-specific nutritional requirements. For eusocial organisms like ants, where only a small fraction of the colony members collect food, increasing evidence demonstrates that strong macronutrient regulation occurs at the colony level. Here, we explored regulation of protein and carbohydrate (p:c) intake in the Rasberry crazy ant, Nylanderia sp. nr. puben. We did this using dry artificial foods (14–42% total macronutrient content) having a range of fixed p:c ratios in a series of choice and no-choice laboratory experiments and used worker mortality to gauge colony-level costs associated with active nutrient regulation. Choice experiments revealed that colonies preferred carbohydrate-rich foods and self-selected a diet having a p:c ratio ~1:2. No-choice experiments demonstrated that food p:c ratio only moderately affected worker food collection behavior, likely because colonies regulated the intake of only the non-limiting nutrients. Absolute worker mortality was generally high, but lowest in colonies feeding on the food having a p:c ratio of 1:2 (the p:c ratio ants self-selected in the choice experiment), although mortality was not significantly affected by food p:c ratio. The self-selected p:c ratio in our study is consistent with that observed in other recent ant nutrient regulation studies. Taken together, the results from these combined studies reveal emerging commonalities among ants in macronutrient regulation strategies, and similarities in foraging behaviors and costs associated with macronutrient regulation. Finally, from a methodological perspective, the high mortality observed in our study, when compared with other recent studies, suggests that ant nutrient regulation studies should be conducted using foods having high moisture and total macronutrient content.     

Summer and fall ants have different physiological responses to food macronutrient content

Seasonally, long-lived animals exhibit changes in behavior and physiology in response to shifts in environmental conditions, including food abundance and nutritional quality. Ants are long-lived arthropods that, at the colony level, experience such seasonal shifts in their food resources. Previously we reported summer- and fall-collected ants practiced distinct food collection behavior and nutrient intake regulation strategies in response to variable food protein and carbohydrate content, despite being reared in the lab under identical environmental conditions and dietary regimes. Seasonally distinct responses were observed for both no-choice and choice dietary experiments. Using data from these same experiments, our objective here is to examine colony and individual-level physiological traits, colony mortality and growth, food processing, and worker lipid mass, and how these traits change in response to variable food protein–carbohydrate content. For both experiments we found that seasonality per se exerted strong effects on colony and individual level traits. Colonies collected in the summer maintained total worker mass despite high mortality. In contrast, colonies collected in the fall lived longer, and accumulated lipids, including when reared on protein-biased diets. Food macronutrient content had mainly transient effects on physiological responses. Extremes in food carbohydrate content however, elicited a compensatory response in summer worker ants, which processed more protein-biased foods and contained elevated lipid levels. Our study, combined with our previously published work, strongly suggests that underlying physiological phenotypes driving behaviors of summer and fall ants are likely fixed seasonally, and change circannually.     

Foraging activity and dietary spectrum of the Argentine ant (Hymenoptera: Formicidae) in invaded natural areas of the northeast Iberian Peninsula

We analyzed the foraging activity and the dietary spectrum of the Argentine ant (Linepithema humile Mayr) and select native ants on cork oaks from Mediterranean open cork oak (Quercus suber) secondary forests. The study areas included invaded and noninvaded zones in close proximity. The Argentine ant's daily foraging activity was correlated to the abiotic factors studied, whereas the seasonal foraging activity was related not only to the variations in the average air temperature, but also to the trophic needs of the colony. Argentine ant workers focused their attention on protein foods during the queens' oviposition periods and during the larvae development phase, and on carbohydrate foods, such as honeydew, when males and workers were hatching. There were no significant differences over the entire year in the quantity of liquid food collected by the Argentine ant workers in comparison with the native ants studied. The solid diet of the Argentine ant on cork oaks is composed of insects, most of which are aphids. Our results have clear applications for control methods based on toxic baits in the invaded natural ecosystems of the Iberian Peninsula.     

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 variation in the collective regulation of foraging by harvester ants

This study investigates variation in collective behavior in a natural population of colonies of the harvester ant, Pogonomyrmex barbatus. Harvester ant colonies regulate foraging activity to adjust to current food availability; the rate at which inactive foragers leave the nest on the next trip depends on the rate at which successful foragers return with food. This study investigates differences among colonies in foraging activity and how these differences are associated with variation among colonies in the regulation of foraging. Colonies differ in the baseline rate at which patrollers leave the nest, without stimulation from returning ants. This baseline rate predicts a colony's foraging activity, suggesting there is a colony-specific activity level that influences how quickly any ant leaves the nest. When a colony's foraging activity is high, the colony is more likely to regulate foraging. Moreover, colonies differ in the propensity to adjust the rate of outgoing foragers to the rate of forager return. Naturally occurring variation in the regulation of foraging may lead to variation in colony survival and reproductive success.     

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.     

Changes in food selection by workers of the pharaoh''s ant, Monomorium pharaonis

The results from simple choice test experiments with laboratory and field colonies of pharaoh's ants have revealed two behavioural traits that influence the way in which foraging worker ants respond to foods. Initially, worker ants from laboratory colonies showed a distinct preference for certain foods (e.g. honey and peanut butter). However, when given only these 'preferred' foods continuously for several weeks, workers showed a marked preference for other foods when offered a choice. This 'satiation' response occurred even though the initial diet was originally highly attractive to foraging workers. In addition, workers show a marked tendency to alternate between carbohydrate foods and protein foods. Thus, workers from colonies fed predominantly on proteins, subsequently showed a marked preference for carbohydrates, and vice versa. The 'satiation' and 'alternation' responses to foods ensure that colonies receive a diet that is both varied and balanced. In addition, these aspects of feeding behaviour have important consequences for the use of food-based baits in control measures against Monomorium pharaonis (L.).     

The farming ant Sericomyrmex amabilis nutritionally manages its fungal symbiont and its social parasite

1. When parasites exploit mutualisms involving food exchange, they can destabilise the partnership with costs to interacting partners. For instance, the ant Sericomyrmex amabilis farms fungal symbionts to produce food, but, in so doing, attracts parasitic Megalomyrmex symmetochus guest ants that infiltrate fungus‐farming ant societies and live with their hosts their entire lives. 2. The present study examined whether host foraging in parasitised colonies shifts towards nutritional requirements of the parasitic guest ants as inferred from the parasite's elemental content (%C, %N, and C:N). 3. Laboratory feeding experiments with nutritionally defined diets indicated that S. amabilis ants harvest protein‐biased substrate, and more total substrate when hosting M. symmetochus relative to when provisioning their fungus gardens and nestmates. 4. Field surveys further showed that parasitised colonies incur reductions in fungus garden nutritional quality and quantity, brood mass, and host worker body condition. And yet these costs appear manageable across growing seasons, as parasitised fungal cultivars appear to provide sufficient nutrition for stable populations of host ants. 5. The approach developed here shows how behavioural strategies for nutrient regulation can extend beyond the needs of the individual to entire fungus‐farming systems, and implies that S. amabilis dynamically adjusts collective foraging strategies when parasitised to enhance long‐term symbiotic stability.     

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.     

The spatial scale of seed collection by harvester ants

Colonies of the seed-eating ant, Pogonomyrmex barbatus, compete with neighboring colonies for foraging areas. In a conflict over foraging area, what is at stake? This depends on how resources are distributed in time and space: if certain regions consistently provide particularly nutritious seed species, or especially abundant seeds, such regions will be of greater value to a colony. During the summer, seeds were taken from returning foragers in colonies located in 4 different vegetation types. There was no relation between the vegetation currently growing in the foraging area, and the species of seeds collected by ants. During the summer, ants collect mostly seeds produced in previous seasons and dispersed by wind and flooding. In 1991, colonies in all vegetation types collected mostly Bouteloua aristidoides; in 1992, Eriastrum diffusum and Plantago patagonica. There was no relation between colony density and numbers of seeds collected. Seed species collected by ants were compared in different colonies, and on different foraging trails within a colony. The results show that seed patches are distributed on the scale of distances between nests, not the smaller scale of different foraging trails of one colony. It appears that colonies are competing for any space in which to search for seeds, not competing for certain regions of consistently high value.     

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.     

Influence of macronutrient imbalance on native ant foraging and interspecific interactions in the field

1. Ants interact with a diversity of organisms. These interactions, coupled with their abundance, cause ants to have ecologically important effects across multiple trophic levels. 2. Empirical study of ant nutritional ecology has led to the prediction that a macronutrient imbalance will affect ant behaviour and interspecific interactions that underlie these broad‐scale effects. Excess carbohydrate relative to protein is predicted to increase ant aggressiveness, predatory tendency and foraging activity, and to decrease collection of hemipteran honeydew and plant nectar. 3. In field experiments conducted in 2009 and 2010, captive colony fragments of a native ant, Formica podzolica (Hymenoptera: Formicidae), were provided with either simulated prey or carbohydrate solution ad libitum. Foraging behaviours and interactions with flowers, myrmecophilous aphids and aphid natural enemies on wild‐grown plants were documented. 4. Strong effects of macronutrient imbalance on foraging manifested quickly and consistently across colonies; in accordance with predictions, prey‐fed foragers collected both honeydew and floral nectar, whereas carbohydrate‐fed ants ceased collecting these resources. Counter to predictions, carbohydrate‐fed ants dramatically lowered their activity levels and did not prey upon aphids. 5. Ants had no effect on aphid enemies in 2009, when the latter were relatively rare, but decreased their abundance in 2010. Despite this protection, the net effect of ants on aphids was negative (measured only in 2009). Prey‐fed ants demonstrated a strong preference for honeydew over floral nectar, thus demonstrating that a macronutrient imbalance may lead to different interactions with similar resources. 6. This study links ant nutrition and community ecology by demonstrating the rapid, asymmetric and multitrophic consequences of nutritionally mediated behaviour.     

Arthropod prey of imported fire ants (Hymenoptera: Formicidae) in Mississippi sweetpotato fields

The red imported fire ants, Solenopsis invicta (Buren), are generally considered pests. They have also been viewed as beneficial predators feeding on other insect pests of various agroecosystems. This study documents the foraging habits of fire ants in a sweetpotato field in Mississippi. Fire ant foraging trails connecting outside colonies to a sweetpotato field were exposed and foraging ants moving out of the field toward the direction of the colony were collected along with the solid food particles they were carrying. The food material was classified as arthropod or plant in origin. The arthropod particles were identified to orders. Fire ant foragers carried more arthropods than plant material. Coleoptera and Homoptera were the most abundant groups preyed upon. These insect orders contain various economically important pests of sweetpotato. Other major hexapod groups included the orders Hemiptera, Diptera and Collembola. The quantity of foraged material varied over the season. No damage to sweetpotato roots could be attributed to fire ant feeding. Imported fire ant foraging may reduce the number of insect pests in sweetpotato fields.     

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.     

The foraging behavior of Japanese macaques Macaca fuscata in a forested enclosure:Effects of nutrient composition,energy and its seasonal variation on the consumption of natural plant foods

In the wild, primate foraging behaviors are related to the diversity and nutritional properties of food, which are affected by seasonal variation. The goal of environmental enrichment is to stimulate captive animals to exhibit similar foraging behavior of their wild counterparts, e.g. To extend foraging time. We conducted a 12-month study on the foraging behavior of Japanese macaques in a semi-naturally forested enclosure to understand how they use both provisioned foods and naturally available plant foods and what are the nutritional criteria of their consumption of natural plants. We recorded time spent feeding on provisioned and natural plant foods and collected the plant parts ingested of their major plant food species monthly, when available.We conducted nutritional analysis (crude protein, crude lipid, neutral detergent fiber-'NDF', ash) and calculated total non-slructural carbohydrate - 'TNC' and total energy of those food items. Monkeys spent 47% of their feeding time foraging on natural plant species. The consumption of plant parts varied significantly across seasons. We found that leaf items were consumed in months when crude protein, crude protein-to-NDF ratio, TNC and total energy were significantly higher and NDF was significantly lower, fruit/nut items in months when crude protein and TNC were significantly higher and crude lipid content was significantly lower, and bark items in months when TNC and total energy were higher and crude lipid content was lower. This preliminary investigation showed that the forested enclosure allowed troop members to more fully express their species typical flexible behavior by challenging them to adjust their foraging behavior to seasonal changes of plant item diversity and nutritional content, also providing the possibility for individuals to nutritionally enhance their diet.     

Getting carbohydrates in ants: damage of the young cereal sprouts by ants of the genus Myrmica Latreille, 1804 (Hymenoptera: Formicidae), to obtain plant sap

Summary

Carbohydrate food is of high importance for survival of ant colonies. Ants are known to use sugary excretions (honeydew) of various insects, nectar of floral and extrafloral nectaries, and even sap of some trees. However, the ability of ants to use sap of herbaceous plants has not been mentioned. This is the first evidence that ants of the genus Myrmica can intentionally ‘cut off’ young cereal sprouts to obtain plant sap. The investigation was carried out in a laboratory in 2018 and 2019 and involved three ant species of the genus Myrmica [12 colonies of M. rubra (Linnaeus, 1758); eight colonies of M. ruginodis Nylander, 1846; and five colonies of M. scabrinodis Nylander, 1846]. First observations were made occasionally in 10 ant colonies during the study of ant–aphid interactions. After three days of carbohydrate starvation, ants were supplied with the plants of wheat infested with aphids of Schizaphis graminum (Rondani, 1852). Within the first day in addition to ordinary trophobiotic relations with aphids, the workers of all the studied colonies demonstrated unexpected behaviour: they ‘cut off’ some sprouts and collected sap of these plants. The experimental investigation in 15 ant colonies of various sizes (about 150, 300 and 500 workers) supplied with the plants infested or non-infested with aphids has shown that getting sap of herbs depends greatly on ant colony needs and available resources. The number of damaged plants was much higher both in the larger colonies of ants and in the absence of aphids. This way of getting carbohydrates allows ants to quickly obtain some extra food needed to maintain colony viability and seems to be one of the mechanisms promoting survival of ants in conditions of acute carbohydrate deficiency. At the same time, ants avoid using plant sap when there are more available alternative carbohydrate resources.     

Functional implications of omnivory for dietary nutrient balance

Captive experiments have shown that many species regulate their macronutrient (i.e. protein, lipid and carbohydrate) intake by selecting complementary food types, but the relationships between foraging strategies in the wild and nutrient regulation remain poorly understood. Using the pine marten as a model species, we collated available data from the literature to investigate effects of seasonal and geographic variation in diet on dietary macronutrient balance. Our analysis showed that despite a high variety of foods comprising the diet, typical of a generalist predator, the macronutrient energy ratios of pine martens were limited to a range of 50–55% of protein, 38–42% of lipids and 5–10% of carbohydrates. This broad annual stabilisation of macronutrient ratios was achieved by using alternative animal foods to compensate for the high fluctuation of particular prey items, and sourcing non‐protein energy (carbohydrates and fats) from plant‐derived foods, particularly fruits. Macronutrient balance varied seasonally, with higher carbohydrate intake in summer–autumn, due to opportunistic fruit consumption, and higher protein intake in winter–spring. In terms of their proportional dietary carbohydrate intake the pine marten's nutritional strategy fell between that of true carnivores (e.g. the wolf) and more omnivorous feeders (e.g. the European badger). However, in terms of energy contributed by protein pine martens are equivalent to obligate carnivores such as the wolf and domesticated cat, and different to some omnivorous carnivores such as the domesticated dog and grizzly bears.     

Mechanisms of carbohydrate‐fuelled ecological dominance in a tropical rainforest canopy‐foraging ant

1. Canopy‐foraging ants have carbohydrate‐rich diets and the stoichiometric excess of carbon may result in energetic allocation decisions that facilitate ecological dominance. 2. If dietary carbohydrates facilitate ecological dominance in canopy ants, then the mechanism for this relationship is unknown. 3. Four hypotheses were posit that may explain how a carbohydrate‐rich diet might facilitate ecological dominance in canopy ants: Aggressive Defense, Metabolic Fuel, Foraging Success, and Prey Acquisition. 4. To assess these hypotheses, experiments were conducted on the canopy‐foraging bullet ant, Paraponera clavata (Fabricius), an omnivorous species that demonstrates high variability in the relative contribution of carbohydrates to the diets of colonies. 5. No support was found for the Aggressive Defense, Metabolic Fuel and Prey Acquisition hypotheses. 6. The Foraging Success hypothesis was supported, as the proportion of nectar in the diet predicted the overall foraging success. 7. It was argued that there is no explicit advantage in the exploitation of nectar over other food resources, other than the fact that it is the most accessible food resource in the rainforest canopy.     

Phorid parasitoids affect foraging activity of Solenopsis richteri under different availability of food in Argentina

1. In Argentina, six species of Pseudacteon parasitoids (Phoridae) attack Solenopsis richteri, one of the two species of South American fire ant that are exotic pests in North America. 2. The presence of these Pseudacteon species significantly reduces the number of ants at food resources in the field, as well as foraging activity generally. 3. Some Pseudacteon not only attack ants walking on trails or at feeding sites, but also at mound entrances, inhibiting workers from leaving to forage. 4. The average size of foraging ants (which prescribes their suitability as hosts) decreased in the presence of phorids. 5. The number of attacking phorids was correlated positively with the number of ants walking towards the food on the trail before the attack. 6. Solenopsis richteri workers responded to manipulations of food size and presence or absence of parasitoids in a risk-adjusting way, i.e. although more foragers were recruited to larger food items, attacking phorids reduced ant foraging activity by the same factor regardless of the size of the food offered. 7. The data suggest that S. richteri colonies juggle the needs to harvest food efficiently, reduce competition, and avoid excess risks from parasitoids in complex ways.     

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.