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.     

Fragmentation or Transportation: Mode of Large-Prey Retrieval in Arboreal and Ground Nesting Ants

Foraging strategy of ant workers has been studied from several aspects, however, the mode of prey retrieval and factors affecting it have been rarely studied to date, even thought it is an important aspect for understanding ant foraging strategy. We investigated the behavioral response against large prey for 44 ant species of 34 genera belonging to eight subfamilies in Japan, Malaysia and Indonesia. Workers of most arboreal ants cut-up large prey at the site of prey capture, and individual workers retrieve the smaller pieces to the nests. In contrast, in most ground-living species, a group of workers retrieve large prey cooperatively without fragmentation. On the ground, parts of the prey item were often robbed by other ant species during this process, while such interference was rare on trees. The significance of the relation between nest site and mode of prey retrieval is discussed.     

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).     

Arboreal Ants Use the “Velcro? Principle” to Capture Very Large Prey

Plant-ants live in a mutualistic association with host plants known as “myrmecophytes” that provide them with a nesting place and sometimes with extra-floral nectar (EFN) and/or food bodies (FBs); the ants can also attend sap-sucking Hemiptera for their honeydew. In return, plant-ants, like most other arboreal ants, protect their host plants from defoliators. To satisfy their nitrogen requirements, however, some have optimized their ability to capture prey in the restricted environment represented by the crowns of trees by using elaborate hunting techniques. In this study, we investigated the predatory behavior of the ant Azteca andreae which is associated with the myrmecophyte Cecropia obtusa. We noted that up to 8350 ant workers per tree hide side-by-side beneath the leaf margins of their host plant with their mandibles open, waiting for insects to alight. The latter are immediately seized by their extremities, and then spread-eagled; nestmates are recruited to help stretch, carve up and transport prey. This group ambush hunting technique is particularly effective when the underside of the leaves is downy, as is the case for C. obtusa. In this case, the hook-shaped claws of the A. andreae workers and the velvet-like structure of the underside of the leaves combine to act like natural Velcro® that is reinforced by the group ambush strategy of the workers, allowing them to capture prey of up to 13,350 times the mean weight of a single worker.     

Defensive behaviour and biological activities of the abdominal secretion in the ant Crematogaster scutellaris (Hymenoptera: Myrmicinae)

Using bioassays, the defensive behaviour of Crematogaster scutellaris and the biological activities of its abdominal secretion were investigated. Beside classical aggressive behaviours such as grips, C. scutellaris workers performed frequent characteristic gaster flexions during interspecific encounters, sometimes tempting to apply their abdominal secretion topically on the enemy. The toxicity of the venom of C. scutellaris to other ants greatly differed among the species tested, some being killed after the topical application of only three droplets, while others were quite resistant to a dose of 90 droplets. All ant species tested were strongly and immediately repelled by a contact between their antennae or mouthparts with the venom of C. scutellaris. Abdominal secretion was never used during intraspecific interference and workers were resistant to a topical application of the venom of their own species. Intraspecific repellency was significant but moderate compared to interspecific one. Workers of C. scutellaris were never seen using their venom during prey capture. In conclusion, the main biological activity of the abdominal secretion of C. scutellaris seems to be its repellency to other ant species. This is supported by field experiments showing that Pheidole pallidula foragers were efficiently repelled at coexploited baits, allowing the monopolization of most prey by C. scutellaris.     

Trophic niche,capture efficiency and venom profiles of six sympatric ant‐eating spider species (Araneae: Zodariidae)

The arms race between specialist predators and their prey has resulted in the evolution of a variety of specific adaptations. In venomous predators, this can include venom composition, particularly if predators are specialized on dangerous prey. Here, we performed an integrative study using six species of highly specialized ant‐eating spiders of the genus Zodarion to investigate their phylogeny, realized trophic niche, efficacy in the capture of various ant species and venom composition. Data on natural diet obtained by next‐generation sequencing and field observations showed that the six Zodarion species exploit different ant species. Their phylogeny, based on mitochondrial and nuclear genes, correlated with the composition of their natural prey, indicating that closely related Zodarion species specialize on similar ant species. Prey‐capture parameters differed among Zodarion species suggesting prey‐specific efficacy. Similarly, the venom profiles of both low and high molecular compounds differed among species. Only the profiles of low molecular compounds were correlated with capture efficacy parameters, suggesting that the venom of Zodarion spiders contains prey‐specific components. Our study suggests that Iberian Zodarion spiders are specialized on particular ant species.     

Mandible strike: the lethal weapon of Odontomachus opaciventris against small prey

In order to study both the hunting efficiency and the flexibility of their predatory behavior, solitary hunters of the trap-jaw ant Odontomachus opaciventris were offered small prey (termites, fruit flies and tenebrionid larvae), presenting different morphological or defensive characteristics. The monomorphic hunters showed a moderately flexible predatory behavior characterized by short capture sequences and a noteworthy efficiency of their mandible strike (76.7-100% of prey retrievals), even when presented with Nasutitermes soldiers. Contrary to most poneromorph ants, antennal palpation of the prey before the attack was always missing, no particular targeted region of the prey's body was preferred, and no 'prudent' posture was ever exhibited. Moreover, stinging was regularly performed on bulky, fast moving fruit flies, very scarcely with sclerotized tenebrionid larvae, but never occurred with Nasutitermes workers or soldiers despite their noxious chemical defense. These results suggest that, whatever the risk linked to potentially dangerous prey, O. opaciventris predatory strategy optimizes venom use giving top priority to the swiftness and strength of the lethal trap-jaw system used by hunters as first strike weapon to subdue rapidly a variety of small prey, ranging from 0.3 to 2 times their own body size and from 0.1 to 2 times their weight. Such risk-prone predatory behavior is likely to be related to the large size of O. opaciventris colonies where the death of a forager might be of lesser vital outcome than in small colony-size 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.     

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.     

Hunting strategy of a generalist ant species proposed as a biological control agent against termites

We studied the hunting behaviour of Myrmicaria opaciventris (Hymenoptera: Formicidae) in order to evaluate if it can be used as a biological control agent against the termites that damage sugarcane plantations. Hunting workers foraged in groups and recruited nestmates at short-range when they encountered large termite soldiers or groups of small termite workers. Differences in prey capture concerned the: (1) means of detection (from a distance or by contact); (2) termite body part seized (small termites seized by the body; large termites by an appendage); (3) percentages of prey abandoned; and (4) use of venom. The sting of the workers is spatulated implying a topical application of the venom on the prey. Large termites were stretched by several workers whose adherence to the substrate is facilitated by well-developed arolia and claws on the legs while others spread venom on the body and carved it up. An adaptation to termite capture was noted with a distribution of tasks between the workers which subdued prey, and those which transported it. In the former case, the workers easily eliminated termite soldiers, successively attacked several termite workers and even captured new individuals while holding the first ones captured between their mandibles before retrieving them all at once. The remaining individuals were retrieved by the transporting workers. Given this particularly effective predatory strategy, we concluded that, under certain conditions, M. opaciventris can be used as a biological control agent against termites.     

Consumption of introduced prey by native predators: Argentine ants and pit-building ant lions

When populations of native predators are subsidized by numerically dominant introduced species, the structure of food webs can be greatly altered. Surprisingly little is known, however, about the general factors that influence whether or not native predators consume introduced species. To learn more about this issue, we examined how native pit-building ant lions (Myrmeleon) are affected by Argentine ant (Linepithema humile) invasions in coastal southern California. Compared to areas without L. humile, invaded areas contained few native ant species and were deficient in medium-sized and large bodied native ants. Based on these differences, we predicted that Argentine ants would negatively affect ant lion larvae. Contrary to this expectation, observational surveys and laboratory growth rate experiments revealed that Myrmeleon were heavier, had longer mandibles, and grew more quickly when their main ant prey were Argentine ants rather than native ants. Moreover, a field transplant experiment indicated that growth rates and pupal weights were not statistically different for larval ant lions reared in invaded areas compared to those reared in uninvaded areas. Argentine ants were also highly susceptible to capture by larval Myrmeleon. The species-level traits that presumably make Argentine ant workers susceptible to capture by larval ant lions—small size and high activity levels—appear to be the same characteristics that make them unsuitable prey for vertebrate predators, such as horned lizards. These results underscore the difficulties in predicting whether or not numerically dominant introduced species serve as prey for native predators.     

Spatial Distribution of Dominant Arboreal Ants in a Malagasy Coastal Rainforest: Gaps and Presence of an Invasive Species

We conducted a survey along three belt transects located at increasing distances from the coast to determine whether a non-random arboreal ant assemblage, such as an ant mosaic, exists in the rainforest on the Masoala Peninsula, Madagascar. In most tropical rainforests, very populous colonies of territorially dominant arboreal ant species defend absolute territories distributed in a mosaic pattern. Among the 29 ant species recorded, only nine had colonies large enough to be considered potentially territorially dominant; the remaining species had smaller colonies and were considered non-dominant. Nevertheless, the null-model analyses used to examine the spatial structure of their assemblages did not reveal the existence of an ant mosaic. Inland, up to 44% of the trees were devoid of dominant arboreal ants, something not reported in other studies. While two Crematogaster species were not associated with one another, Brachymyrmex cordemoyi was positively associated with Technomyrmex albipes, which is considered an invasive species—a non-indigenous species that has an adverse ecological effect on the habitats it invades. The latter two species and Crematogaster ranavalonae were mutually exclusive. On the other hand, all of the trees in the coastal transect and at least 4 km of coast were occupied by T. albipes, and were interconnected by columns of workers. Technomyrmex albipes workers collected from different trees did not attack each other during confrontation tests, indicating that this species has formed a supercolony along the coast. Yet interspecific aggressiveness did occur between T. albipes and Crematogaster ranavalonae, a native species which is likely territorially dominant based on our intraspecific confrontation tests. These results suggest that the Masoala rainforest is threatened by a potential invasion by T. albipes, and that the penetration of this species further inland might be facilitated by the low density of native, territorially dominant arboreal ants normally able to limit its progression.     

Alien and native tree species having extrafloral nectaries as favorite hunting area for arboreal endemic Philippine tiger beetles (Coleoptera: Cicindelidae) in human‐disturbed habitat in Lanao del Sur Province,Mindanao, Philippines

To document a relation between abundance of arboreal, predatory tiger beetles, their ant prey, and extrafloral nectaries attracting the ants, we gathered data from more than 10 species of native and introduced trees and large, tree‐like perennial plants in Lanao del Sur Province, Mindanao, Philippines. All specimens of tiger beetles (two Tricondyla and two Neocollyris species, all endemic to the country) were noted on five tree species characterized by presence of extrafloral nectaries, including three alien/invasive and two native ones. Invasive Spathodea campanulata and native Hibiscus tiliaceus were the most inhabited ones (respectively, 56% and 19% of beetles). Presence of tiger beetles on these trees most probably depends on high abundance of ants, which are typical prey for arboreal Cicindelidae, while occurrence of ants can result from presence of extrafloral nectaries on different parts of the plants. This suggests a new mutualistic insect–plant interaction between native and invasive species.     

Sugary food robbing in ants: a case of temporal cleptobiosis

This study reports new information on interactions between Ectatomma tuberculatum (Ponerinae) and Crematogaster limata parabiotica (Myrmicinae). Workers of these sympatric arboreal ant species forage on the same pioneer trees. Diurnally, Ectatomma preyed on Crematogaster workers that avoided overt aggression by respecting a 'safe distance'. At night, Crematogaster initiated raids within the Ectatomma nests that they apparently left with their abdomen empty, then remained near the nest entrances where they successfully intercepted 75.2% of the returning Ectatomma foragers (N = 322). Certain intercepted workers rapidly resumed their return trip. Others (39.1%) were stopped, explored and licked during a long time by the Crematogaster. Most of them were carrying between their mandibles a droplet of liquid food that was stolen. This relationship, that appears to be a typical case of interspecific cleptobiosis, whose expression varies during the daytime, demonstrates for the first time sugary-food robbing, instead of prey robbing, in ants.     

The arboreal ants of a Neotropical rain forest show high species density and comprise one third of the ant fauna

In tropical rain forests, the ant community can be divided into ground and arboreal faunas. Here, we report a thorough sampling of the arboreal ant fauna of La Selva Biological Station, a Neotropical rain forest site. Forty-five canopy fogging samples were centered around large trees. Individual samples harbored an average of 35 ant species, with up to 55 species in a single sample. The fogging samples yielded 163 observed species total, out of a statistically estimated 199 species. We found no relationship between within-sample ant richness and focal tree species, nor were the ant faunas of nearby trees more similar to each other than the faunas of widely spaced trees. Species density was high, and beta diversity was low: A single column of vegetation typically harbors at least a fifth of the entire arboreal ant fauna. Considering the entire fauna, based on 23,326 species occurrence records using a wide variety of collecting methods, 182 of 539 observed species (196 of 605, estimated statistically) were entirely arboreal. The arboreal ant fauna is thus about a third of the total La Selva ant fauna, a robust result because inventory completeness was similar for ground and arboreal ants. The taxonomic history of discovery of the species that make up the La Selva fauna reveals no disproportionately large pool of undiscovered ant species in the canopy. The "last biotic frontier" for tropical ants has been the rotten wood, leaf litter, and soil of the forest floor.     

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.     

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).     

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'.     

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.     

Predatory behavior of a seed-eating ant:Brachyponera senaarensis

The great flexibility of the feeding strategies exhibited by the ponerine ant Brachyponera senaarensis (Mayr) allows it to exploit either seeds or animal prey items as food resources. Predation is generally limited to small prey and is very similar to scavenging behavior. In laboratory conditions, the predatory behavior of B. senaarensis is not different in structure from that known in other carnivorous ants species. The workers forage individually and return to the nest using a series of cues involving light, a chemical graduated marking system near the nest entrance, and memory. During nest-moving, recruitment by tandem running was observed. However, in colonies where the food supply is regular, workers that discover food do not recruit nestmates, but make repeated trips between the nest and the food source. On the contrary, in starved colonies, the introduction of prey may produce a massive exit of foragers, corresponding to a primitive form of mass recruitment similar to that observed in some other ant species.