The Effect of Food Size and Dispersion Pattern on Retrieval Rate by the Argentine Ant, Linepithema humile (Hymenoptera: Formicidae)

Food acquisition in central-place foraging animals demands efficient detection and retrieval of resources. Most ant species rely on a mass recruitment foraging strategy, which requires that some potential foragers remain at the nest where they can be recruited to food once resources are found. Because this strategy reduces the number of workers initially looking for food, it may reduce the food detection rate while increasing the postdiscovery food retrieval rate. In previous studies this tradeoff has been analyzed by computer simulation and mathematical models. Both kinds of models show that food acquisition rate is greatly influenced by food distribution and resource patch size: as food is condensed into fewer patches, the maximal acquisition rate is achieved by a shift to fewer initial searchers and more potential recruits. In general, these models show that a mass recruitment strategy is most effective when resources are clumped. We tested this prediction in two experiments by letting laboratory colonies of the Argentine ant (Linepithema humile) forage for resources placed in different distributions. When all prey were small, retrieval rate increased with increasing resource patch size, in support of foraging models. When prey were large, however, the mass of prey returned to the colony over time was much lower than when prey were small and widely distributed. As more ants reached a large prey item, the distance the prey item was transported decreased due to a greater emphasis on feeding rather than transport. Because Argentine ants can transport more biomass externally than they can ingest, food retrieval that depends only on ingestion can depress the biomass retrieval rate. Thus, our results generally support theoretical foraging models, but we show how prey size, through differential prey-handling behavior, can produce an outcome greatly different from that predicted only on the distribution of resources.     

Prey capture behavior in an arboreal African ponerine ant

I studied the predatory behavior of Platythyrea conradti, an arboreal ponerine ant, whereas most species in this subfamily are ground-dwelling. The workers, which hunt solitarily only around dusk, are able to capture a wide range of prey, including termites and agile, nocturnal insects as well as diurnal insects that are inactive at that moment of the Nyctemeron, resting on tree branches or under leaves. Prey are captured very rapidly, and the antennal palpation used by ground-dwelling ponerine species is reduced to a simple contact; stinging occurs immediately thereafter. The venom has an instant, violent effect as even large prey (up to 30 times the weight of a worker) never struggled after being stung. Only small prey are not stung. Workers retrieve their prey, even large items, singly. To capture termite workers and soldiers defending their nest entrances, ant workers crouch and fold their antennae backward. In their role as guards, the termites face the crouching ants and end up by rolling onto their backs, their legs batting the air. This is likely due to volatile secretions produced by the ants' mandibular gland. The same behavior is used against competing ants, including territorially-dominant arboreal species that retreat further and further away, so that the P. conradti finally drive them from large, sugary food sources.     

Species‐level predation network uncovers high prey specificity in a Neotropical army ant community

Army ants are among the top arthropod predators and considered keystone species in tropical ecosystems. During daily mass raids with many thousand workers, army ants hunt live prey, likely exerting strong top‐down control on prey species. Many tropical sites exhibit a high army ant species diversity (>20 species), suggesting that sympatric species partition the available prey niches. However, whether and to what extent this is achieved has not been intensively studied yet. We therefore conducted a large‐scale diet survey of a community of surface‐raiding army ants at La Selva Biological Station in Costa Rica. We systematically collected 3,262 prey items from eleven army ant species (genera Eciton, Nomamyrmex and Neivamyrmex). Prey items were classified as ant prey or non‐ant prey. The prey nearly exclusively consisted of other ants (98%), and most booty was ant brood (87%). Using morphological characters and DNA barcoding, we identified a total of 1,103 ant prey specimens to the species level. One hundred twenty‐nine ant species were detected among the army ant prey, representing about 30% of the known local ant diversity. Using weighted bipartite network analyses, we show that prey specialization in army ants is unexpectedly high and prey niche overlap very small. Besides food niche differentiation, we uncovered a spatiotemporal niche differentiation in army ant raid activity. We discuss competition‐driven multidimensional niche differentiation and predator–prey arms races as possible mechanisms underlying prey specialization in army ants. By combining systematic prey sampling with species‐level prey identification and network analyses, our integrative approach can guide future research by portraying how predator–prey interactions in complex communities can be reliably studied, even in cases where morphological prey identification is infeasible.     

Recruitment Behavior During Foraging in the Neotropical Ant Gnamptogenys moelleri (Formicidae: Ponerinae): Does the Type of Food Matter?

Gnamptogenys moelleri nests in bromeliads and feeds on an array of food items, including dead and live animals, and nectar. Field data in Brazilian forests indicate that G. moelleri hunts solitarily, while retrieving is performed both by solitary workers for small items, or by a group of recruited workers for large items. This flexible foraging strategy was investigated in the laboratory through a series of experiments to assess the context in which recruitment is elicited. Three types of food were used: 50% honey solution, large insect prey, and cluster of small insects. For all food types the first encounter by a scout resulted in increased numbers of ants leaving the nest and finding the food in the arena. After finding liquid food or large prey, the forager returns to the nest and transmits information to nestmates about food location on the substrate. The successful scout repeatedly taps the sting on the ground, and recruited ants collectively retrieve the large insect to the nest. On the other hand, there is no transmission of information to nestmates about the location of small clumped prey, although the returning scout induces nestmates to leave the nest and hunt. Because foraging in G. moelleri is restricted mostly to the nest bromeliad, and small worker size (0.5 cm) precludes capturing large prey solitarily, recruitment behavior widens the spectrum of food items consumed by this ant species. Although recruitment behavior in ponerines has already been reported to vary with the type and size of a food source, this study also shows that the transmission of information about food location depends on the type of food found (large prey or liquid food versus cluster of small prey).     

Prey concealment and spatiotemporal patrolling behaviour of the Indian tree ant Oecophylla smaragdina (Fabricius)

Summary: Oecophylla smaragdina workers conceal prey with leaves and twigs on discovering large prey on their ground territory. The prey concealment behaviour occurs concurrently with prey capture and killing. Ants were observed conducting spatiotemporal patrolling around large prey. The garden lizard Calotes versicolor and at least two predatory ant species were found to forage in the same area as O. smaragdina. It is suggested that the prey concealment behaviour of Oecophylla may be a strategy to prevent prey detection by vertebrate predators which use vision to prey on similar large prey species.     

Prey weight and overwhelming difficulty impact the choice of retrieval strategy in the Neotropical ant <Emphasis Type="Italic">Gnamptogenys sulcata</Emphasis> (F. Smith)

By regulating both the choice between solitary or collective retrieval and the level of nestmate investment for heavy prey transport, individual hunters of the ant Gnamptogenys sulcata are able to increase the foraging efficiency of the whole society, despite the extremely small size of this elite group (1 –4 hunters per colony). Their predatory behavior changes according to the static (weight, size, shape) and dynamic (mobility, escape behaviors) characteristics of the prey. The behavioral sequences resemble those of other poneromorph ants, but palpation after the approach phase is absent, probably because of the swiftness of the attack. Hunting is always performed by solitary workers but, independently of prey type and mobility, small, light prey trigger solitary retrieval whereas large, heavy prey trigger collective retrieval. For intermediate prey weights (8.5 to 21 times the hunter’s weight), some variability in the strategy choice is encountered. Both static and dynamic factors are involved in assessing the difficulty of handling living prey as shown by the number of stings delivered by the hunter for prey immobilization. However, the lack of any stinging against already dead prey indicates that these factors must be uncoupled when selecting retrieval strategies. Prey weight has a major role in this choice. For a given prey size always triggering solitary retrieval, a two-fold increase in prey weight is sufficient to trigger collective transport. Conversely, for a given prey size always triggering collective retrieval, a decrease in prey weight can trigger solitary transport. Moreover, presenting hunters with prey of “infinite” weight triggers several waves of recruitment, so that the number of recruited workers compensates for the “apparent” prey weight. Despite the inability to finely match the number of recruits to the weight of retrieved prey, recruitment strategies adopted by G. sulcata may reflect an evolutionary primitive step towards the finely graded recruitment behavior observed in closely related poneromorph species. Received 5 February 2007; revised 25 May 2007; accepted 15 June 2007.     

Predatory Behavior in Dominant Arboreal Ant Species: The Case of Crematogaster sp. (Hymenoptera: Formicidae)

Crematogaster sp. is a dominant arboreal ant species that captures and retrieves very large prey. Hunting workers forage collectively thanks to short-range recruitment. They detect prey by contact, then rapidly attack, seizing small prey by the body and large prey by a leg. In this study, almost all the active prey were spread-eagled by several workers, even when small enough to permit a single worker to easily master them. While certain workers spread-eagled the prey, others deposited venom on the prey body using their spatulated sting (topical action of the venom). The well-developed arolia on the pretarsus of workers' legs have crucial importance for the success of prey capture (spread-eagling) and transport in an arboreal habitat. These results are compared with those known for other arboreal-dwelling generalist predator ant species.     

Steep Decline and Cessation in Seed Dispersal by Myrmica rubra Ants

Myrmecochorous diaspores bear a nutrient-rich appendage, the elaiosome, attractive to ant workers that retrieve them into the nest, detach the elaiosome and reject the seed intact. While this interaction is beneficial for the plant partner by ensuring its seed dispersal, elaiosome consumption has various effects −positive, negative or none − on ants’ demography and survival, depending on both the ant/plant species involved. In this context, the contribution of ants to seed dispersal strongly varies according to the ant/plant pairs considered. In this paper, we investigate whether the dynamics of myrmecochory also vary on a temporal scale, for a given pair of partners: Myrmica rubra ants and Viola odorata seeds. During their first encounter with seeds, ants collect all the diaspores and eat the majority of elaiosomes. Both the harvesting effort and the elaiosome consumption decline when seeds are offered on the next week and completely cease for the following weeks. This is related to a decrease in the number of foragers reaching the food source, as well as to a reduced probability for an ant contacting a seed to retrieve it. Seed retrieval is not reactivated after seven weeks without any encounter with V. odorata seeds. By contrast, naive ant colonies only fed with fruit flies do not show a decline of prey harvesting of which the speed of retrieval even increases over the successive weeks. Myrmecochory may thus be labile at the scale of a fruiting season due to the ability of ants to steeply tune and cease for several months the harvesting of these seemingly poorly rewarding items and to maintain cessation of seed exploitation. The present study emphasizes the importance of a long-lasting follow up of the myrmecochory process, to assess the stability of this ant-plant partnership and to identify mechanisms of adaptive harvesting in ants.     

Essential and alternative prey in a ponerine ant: variations according to the colony life cycle

We studied the prey specialization of Plectroctena minor, a ponerine ant known to capture mostly millipedes. We compared the prey spectrum of the hunting workers from large colonies with that of the founding queens. The hunting workers captured all kinds of tested prey, but hunted mostly millipedes. Founding queens, which avoided relatively large prey, including the millipedes tested, captured mostly isopods under experimental conditions. We also verified that the presence of millipedes in the diet of the larvae of large colonies was necessary for the production of winged females and strongly enhanced the production of workers, permitting us to assert that P. minor is a predatory species specialized in the capture of millipedes. In contrast, the presence of millipedes had no impact on the production of males. We thus assert that millipedes constitute the 'essential prey' of P. minor, while other arthropod taxa are therefore 'alternative prey'.     

The ecology and feeding habits of the arboreal trap-jawed ant Daceton armigerum

Here we show that Daceton armigerum, an arboreal myrmicine ant whose workers are equipped with hypertrophied trap-jaw mandibles, is characterized by a set of unexpected biological traits including colony size, aggressiveness, trophobiosis and hunting behavior. The size of one colony has been evaluated at ca. 952,000 individuals. Intra- and interspecific aggressiveness were tested and an equiprobable null model used to show how D. armigerum colonies react vis-à-vis other arboreal ant species with large colonies; it happens that D. armigerum can share trees with certain of these species. As they hunt by sight, workers occupy their hunting areas only during the daytime, but stay on chemical trails between nests at night so that the center of their home range is occupied 24 hours a day. Workers tend different Hemiptera taxa (i.e., Coccidae, Pseudococcidae, Membracidae and Aethalionidae). Through group-hunting, short-range recruitment and spread-eagling prey, workers can capture a wide range of prey (up to 94.12 times the mean weight of foraging workers).     

Army ant behaviour in the poneromorph hunting ant Onychomyrmex hedleyi Emery (Hymenoptera: Formicidae; Amblyoponinae)

Onychomyrmex belongs to the phylogenetically basal ant tribe Amblyoponini but shows prototypical army ant behaviours, i.e. group predation and nomadism. In order to investigate these behaviours, Onychomyrmex hedleyi was observed in the field and in laboratory experiments. Workers of O. hedleyi would frequently hunt centipedes but rarely social insects. Workers did not dismember the victims but recruited the colony mates to conduct group retrieval. If the prey were too large or too heavy to retrieve, the entire colony moved from the bivouac site to the prey site. Although foraging on the forest floor, a colony repeated the extension and withdrawal of a raiding column, which was up to 80 cm long (mean ± SD, 41.6 ± 18.5 cm). Colonies were nomadic and the relocation distance was up to 150 cm (mean ± SD, 74.4 ± 45.0 cm). Retinues guarded a queen who moved to a new bivouac site in the early phase of relocation. Colonies were found to stay at a site statistically longer if they had come from a more distant site, and were also observed to move to a more distant site if they had spent a longer time at a particular bivouac site. The consecutive migrations did not show significant directionality.     

Interspecific displacement mechanisms by the invasive little fire ant <Emphasis Type="Italic">Wasmannia auropunctata</Emphasis>

Competition between invasive species and native ones in the new environment was found to be significant and to affect both animal and plant species. Invasive ants are notorious for displacing local ant species through competition. Competitive displacement of native species can occur through interference and or resource competition. However, for invasive ants, little is known about the relative importance of competitive displacement. We studied competitive interactions of the little fire ant, Wasmannia auropunctata, one of the most destructive invasive ant species, with two other ant species, Monomorium subopacum and Pheidole teneriffana. We compared the species’ foraging behavior and studied their aggressive interactions around food baits for the short (2 h) and long (21 days) term in the laboratory. Surprisingly we found that in short term experiments W. auropunctata had the poorest foraging abilities of the three species studied: it took the workers the longest to locate the bait and retrieve it; in addition they retrieved the lowest amount of food. When both W. auropunctata and M. subopacum were foraging the same bait, in the short term competition experiment, W. auropunctata workers did not defend the bait, and ceased foraging when encountered with competition. The long-term experiments revealed that W. auropunctata had the advantage in aggressive interactions over time; they eliminated seven of nine M. subopacum’s nests while consuming some of the workers and brood. According to our laboratory studies, W. auropunctata cannot be considered an extirpator species, unless it has a substantial numerical advantage, in contrast with previous assumptions. Otherwise it may behave as an insinuator species, i.e. the workers do not initiate aggression and by staying undetected they can continue foraging adjacent to dominant species.     

The hunting behavior of the African ponerine ant Pachycondyla pachyderma

The hunting behavior of the African ponerine ant Pachycondyla pachyderma, a semi-specialized centipede predator, appears well adapted to this kind of prey and shows a graded complexity according to the difficulty it has in overwhelming prey. Small prey (5-to-8-mm-long termites) were detected by contact and seized by the thorax while larger prey (≥30-mm-long centipedes) were frequently detected from a distance and seized by the anterior-most part of their body. Termites and 30-mm-long lithobiomorph centipedes were not always stung, whereas stinging and even repeated stinging was needed for 50-mm-long geophilomorphs and scolopendromorphs. Moreover, overwhelming wide and heavy scolopendromorphs, which have better defensive abilities, involved the use of additional behaviors allowing the workers to capture them safely: venom spreading, and a peculiar stinging posture, the “fatal embrace”. Here the workers seize scolopendromorphs by an antenna or by one of their first legs, wrap themselves around the prey while maintaining their grip with their mandibles and legs, and slowly inject venom into the prey's ventral surface. Workers retrieve small prey solitarily while, for large geophilomorphs and scolopendromorphs, nestmates can be recruited at short range or even at long range through tandem running.     

Foraging decisions of individual workers vary with colony size in the greenhead ant Rhytidoponera metallica (Formicidae, Ectatomminae)

Summary. The ability of worker ants to adapt their behaviour depending on the social environment of the colony is imperative for colony growth and survival. In this study we use the greenhead ant Rhytidoponera metallica to test for a relationship between colony size and foraging behaviour. We controlled for possible confounding ontogenetic and age effects by splitting large colonies into small and large colony fragments. Large and small colonies differed in worker number but not worker relatedness or worker/brood ratios. Differences in foraging activity were tested in the context of single foraging cycles with and without the opportunity to retrieve food. We found that workers from large colonies foraged for longer distances and spent more time outside the nest than foragers from small colonies. However, foragers from large and small colonies retrieved the first prey item they contacted, irrespective of prey size. Our results show that in R. metallica, foraging decisions made outside the nest by individual workers are related to the size of their colony.Received 23 March 2004; revised 3 June 2004; accepted 4 June 2004.     

Convergent evolution,superefficient teams and tempo in Old and New World army ants

Swarm raiding army ants, with hundreds of thousands or millions of workers per colony, have evolved convergently in the Old World and New World tropics. Here we demonstrate for the first time, to our knowledge, superefficient foraging teams in Old World army ants and we compare them quantitatively with such teams in New World army ants. Colonies of Dorylus wilverthi in the Old World and Eciton burchelli in the New World retrieve almost identical sizes of prey item and the overall size range of their workers is very similar. However, 98% of D. wilverthi workers are within the size range of the smallest 25% of E. burchelli workers. In E. burchelli larger workers specialize in prey retrieval, whereas in D. wilverthi workers form many more teams than in E. burchelli. Such teams compensate for the relative rarity of larger workers in Dorylus. The proportions of prey items retrieved by teams in Dorylus and Eciton are 39% and 5%, respectively. The percentages of all prey biomass retrieved by teams in Dorylus and Eciton are 64% and 13%, respectively. Working either as single porters or teams, Dorylus carry more per unit ant weight than do Eciton, but Eciton are swifter. However, these different ergonomic factors counterbalance one another, so that performance at the colony level is remarkably, although by no means completely, similar between the Old and New World species. The remaining differences are attributable to adaptations in worker and colony tempo associated with the recovery dynamics of their prey populations. Our comparative analysis provides a unique perspective on worker-level and colony-level adaptations and is a special test of the theory of worker caste distributions.     

Army ant prey availability and consumption by chimpanzees (Pan troglodytes vellerosus) at Gashaka (Nigeria)

Army ant predation by chimpanzees has been studied as an intriguing example of tool use and a possible case of cultural variation. However, the importance of army ant prey in chimpanzee diet and feeding ecology is still only poorly understood. We studied the availability and consumption of army ants in a population of the chimpanzee subspecies Pan troglodytes vellerosus in Nigeria. Army ants were collected from nests and trails (workers) and near artificial light sources (males). Three potential prey species were found: Dorylus rufescens, Dorylus gerstaeckeri and Dorylus kohli . Dorylus rufescens was by far more abundant than the other two species. Only remains from D. rufescens were present in chimpanzee faeces. This is the first report of consumption of this ant species by chimpanzees. However, because of the low availability of the other two species, it is unclear whether this pattern reflects a preference for D. rufescens . Although D. rufescens ' availability varied with weather conditions, the occurrence as well as the absolute and relative numbers of Dorylus fragments in faeces did not. This finding, together with the considerable difficulties encountered by human observers in their efforts to locate nests by following trails, suggests that the chimpanzees in this population do not harvest army ants from trails and do not use trails to locate nests. The overall occurrence of army ant fragments in 42.3% of all faecal samples is the highest ever recorded in any chimpanzee population. This indicates that in this chimpanzee population, army ant prey is not a fallback during periods of sparse availability of plant food, but quantitatively important throughout the year. Future studies will be needed to clarify which cues and strategies chimpanzees use to locate army ant nests and to assess the role of myrmecophagy with respect to macro- or micronutrient demands.     

Prey spectra of two swarm-raiding army ant species in East Africa

Although swarm-raiding army ants are considered keystone predators in tropical forest ecosystems, information on the prey spectra of most species is based on anecdotal reports and not on systematic studies with extensive sampling. We analysed prey samples of 18 colonies of the two afrotropical species Dorylus ( Anomma ) molestus and Dorylus ( Anomma ) wilverthi (4289 prey items in total) to examine the prey composition variation within and between species and to determine the best methodology to obtain reliable prey spectrum estimates for a given species, site and season. Variation in prey composition was substantial for D. molestus even within a single site and season, with the biomass proportion of the most important prey type differing by a factor of 12. Conclusions from studies using small samples sizes may thus be misleading. We demonstrate that the method of assessing prey spectra in terms of relative prey item numbers often produces biased results, and therefore recommend relative prey biomass as the more useful parameter. The near absence of earthworms, which always constituted a substantial part of the D. molestus prey, in the diet of D. wilverthi is interpreted to result from subtle differences in swarm-raiding behaviour between these two species, but could alternatively also be due to low availability. Similar studies recording prey composition as biomass proportions and analysing large samples sizes from many colonies are needed to understand the effect of army ant swarm raids on invertebrate communities in afrotropical forests.     

The predatory behaviour of a tramp ant species in its native range

Workers of the pest ant Paratrechina longicornis participate in a type of group hunting. Each individual forages with its long antennae wide open and moves quickly (6.3 cm/s) along an erratic path surrounded by nestmates behaving in the same way and within range of a recruiting pheromone. They detect prey by contact with successful workers singly capturing and retrieving small prey and seizing larger ones by an appendage. Then they recruit nestmates at short-range; all together they spread-eagle the prey and retrieve them whole.     

Patterns in foraging and nesting ecology in the neotropical ant, <Emphasis Type="Italic">Gnamptogenys moelleri</Emphasis> (Formicidae,Ponerinae)

Summary This study provides quantitative field data on the natural history and foraging behaviour of the Neotropical bromeliad-nesting ant Gnamptogenys moelleri (Ponerinae) in a sandy plain forest in Southeast Brazil. The ant nested on different bromeliad species and the nests were more frequently found in bigger bromeliads. The species used a wide array of invertebrates in its diet, hunting for live prey and scavenging the majority of the items from dead animals. The food items varied greatly in size (1 to 26 mm). Hunting was always performed by solitary workers. Retrieving was performed by solitary workers (small items), or by a group of 3 to 12 workers recruited to the food source (large items). Almost all G. moelleri foraging activity was restricted to the nest bromeliad. In the warm period more ants left the nest to forage, and foraging trips achieved greater distances compared to the cool season. Trap data revealed that overall availability of arthropod prey is higher in the summer than in the winter. The opportunism in nest site use and in foraging behaviour, the small foraging area, as well as the seasonal differences in foraging activity are discussed and compared with other tropical ants.Received 30 May 2003; revised 22 September 2003; accepted 3 October 2003.     

Changes in the speed of ants as a result of aggressive interactions

Subordinate ant species utilize different tactics to reduce competition with the stronger, larger and more aggressive individuals of a dominant species. In our experimental study, we assessed the behavioral response of individual workers of 4 subordinate ant species during their co‐occurrence with workers of a single dominant species. Contrary to most classical experiments focused on aggressive interactions, we assessed workers’ speed as a crucial factor in the outcome of co‐occurrence. Generally, there was a large intraspecific variation in the speed of the studied species—each had slow and fast individuals. Workers of all studied species moved faster just after interaction, suggesting that contact between 2 hostile workers is a stressful stimulus, generating a behavioral reaction of increasing speed. Also, the number of aggressive contacts experienced by a given individual positively affected its speed. Moreover, workers which were fast when exploring territory were also fast after interspecific interactions. The duration of aggression was significantly reduced by the speed and body size of a subordinate species worker—the more quickly a worker reacted and bigger it was, the shorter was the time of cumulative aggression. To our knowledge, this is the first study of this type to be conducted on ants and we conclude that speed is an overlooked and important characteristic of species and also individuals, therefore it should be considered as a driver of patterns of co‐occurrence in ant assemblages.