Distance effects on resource profitability and recruitment in the giant tropical ant,Paraponera clavata

Summary We examine how cost and benefit components of resource profitability affect recruitment in the giant tropical ant, Paraponera clavata. To vary resource profitability, we changed the quantity of artificial nectar baits presented to foragers and the distance of nectar baits from the nest. Both distance to and amount of resource affected quantitative aspects of recruitment. At increased distances foragers were less likely to recruit, and fewer workers were recruited to the resource area. The amount of nectar affected the tendency of foragers to recruit, but had no effect on the number of ants recruited. Variation in resource distance was also associated with qualitative changes in recruitment strategy. Foragers at distant sites recruited from the canopy rather than from the nest, and often transferred nectar to other workers for transport to the nest. Nectar transfer and extra-nidal recruitment significantly reduced the time required for resource collection. It may also have increased the ability of workers to specialize in specific foraging tasks. A portion of the colony's foraging force specialized spatially by remaining in distant foraging areas without returning to the nest. The flexible recruitment system of P. clavata increases colonial net energetic gain rates by concentrating foraging effort on resources yielding the highest net energetic rewards, and increases the competitive abilities of individual colonies at resource sites by decreasing collection times.     

Forest type affects prey foraging of saddleback tamarins,Saguinus nigrifrons

Callitrichids can persist in secondary forests where they may benefit from elevated prey abundance. However, how tamarins forage for prey in secondary forest compared to primary forest has not been examined. Using scan and focal sampling, we compared prey foraging and capture success of two groups of Saguinus nigrifrons in north-eastern Peru: one ranging in primary forest, the other with access to a 10-year-old anthropogenic secondary forest. There was a trend for more prey search in the secondary forest, but prey feeding, capture success and size were lower compared to the primary forest. Tamarins avoided the forest floor, used vertical supports less often and searched on a lower variety of substrates in the secondary forest. In the secondary forest, tamarins did not capture flushed prey, which make up a substantial part of the total prey captures biomass in primary forests. Reduced prey capture success is unlikely to reflect reduced prey availability, since more Orthoptera were found in secondary forest through ultrasonic surveys. Therefore, the prey search activity of S. nigrifrons in young secondary forests seemed rather opportunistic, presumably influenced by altered predation patterns, vegetation structure, as well as prey diversity.     

Flying Foxes Prefer to Forage in Farmland in a Tropical Dry Forest Landscape Mosaic in Fiji

To test flying fox adaptations to a habitat mosaic with extreme deforestation, the abundance, habitat choice and feeding behavior of the Pacific flying fox, Pteropus tonganus, were investigated across 16 islands of the Yasawa archipelago, Fiji. The habitat mosaic is formed by 4.3 percent tropical dry forest and 3.3 percent farmland, leaving exotic grasslands and stands of Leucaena leucocephala to overrun the vast majority of land. Pteropus tonganus abundance was high (5757 bats) despite deforestation and hunting. Roosting sites were restricted to native forest fragments. Grasslands and stands of L. leucocephala were completely void of bats at all times. The mean foraging density in farmland was four times higher than in forests and foraging competition was routinely observed in farmland but was extremely rare in forests. The author suggests that during the study, extensive foraging in farmland was supporting the high P. tonganus population. Additionally, the preferential foraging in farmland was responsible for the low foraging densities within forests and dramatically less intraspecies competition for forest resources. Further research is needed on seed dispersal within forests and to test for seasonal variations in bat abundance and feeding.     

The Dynamics of Foraging Trails in the Tropical Arboreal Ant Cephalotes goniodontus

The foraging behavior of the arboreal turtle ant, Cephalotes goniodontus, was studied in the tropical dry forest of western Mexico. The ants collected mostly plant-derived food, including nectar and fluids collected from the edges of wounds on leaves, as well as caterpillar frass and lichen. Foraging trails are on small pieces of ephemeral vegetation, and persist in exactly the same place for 4–8 days, indicating that food sources may be used until they are depleted. The species is polydomous, occupying many nests which are abandoned cavities or ends of broken branches in dead wood. Foraging trails extend from trees with nests to trees with food sources. Observations of marked individuals show that each trail is travelled by a distinct group of foragers. This makes the entire foraging circuit more resilient if a path becomes impassable, since foraging in one trail can continue while a different group of ants forms a new trail. The colony’s trails move around the forest from month to month; from one year to the next, only one colony out of five was found in the same location. There is continual searching in the vicinity of trails: ants recruited to bait within 3 bifurcations of a main foraging trail within 4 hours. When bait was offered on one trail, to which ants recruited, foraging activity increased on a different trail, with no bait, connected to the same nest. This suggests that the allocation of foragers to different trails is regulated by interactions at the nest.     

ANT DISPERSAL OF CALATHEA (MARANTACEAE) SEEDS BY CARNIVOROUS PONERINES (FORMICIDAE) IN A TROPICAL RAIN FOREST

Two tropical herbaceous plant species, Calathea microcephala and Calathea ovandensis, exhibit morphological adaptations for seed dispersal by ants. In the field, 21 ant species are attracted to the arils of the seeds. Previously known as predatory carnivores, the ponerine ants Odontomachus laticeps, O. minutus, Pachycondyla harpax, and P. apicalis carry the seeds toward their nests, behaving as though they were carrying prey. These ants remove the aril in the nest, and seeds without arils germinate more readily than seeds with arils. Ant foraging distances can account for seedling distributions. Myrmecochory may be far more common in tropical rain forests than has been previously suspected.     

Predation of army ants by Toppin’s titi monkey, <Emphasis Type="Italic">Plecturocebus toppini</Emphasis> Thomas 1914 (Primates: Pitheciidae), in an urban forest fragment in eastern Acre,Brazil

The predation of army ants (Eciton rapax) was recorded during an observational study of the feeding behavior of a group of titi monkeys (Plecturocebus toppini) in an urban fragment of forest in Acre, Brazil. During one observed event, the group’s adult female used its tail to retrieve ants, a type of behavior not observed previously in this genus. All incidents of on-forest floor foraging occurred during the dry season, when fruit was least abundant in the forest, while on other occasions, the ants were captured from tree branches and leaves. The observation of predation and ground-level foraging recorded in this study reinforce the adaptive capacity of P. toppini for survival in fragmented forests, and this was also the first record of the predation of army ants by this titi species.     

Placing an Omnivore in a Complex Food Web: Dietary Contributions to Adult Biomass of an Ant

Workers of Paraponera clavata, a common Neotropical ant, collect both nectar and insect prey. Previous reports show that nectar accounts for up to 90 percent of the ants’ food loads, while calculations suggest that nectar contributes only 10 percent of colonies’ energy supply. We assessed the trophic source of carbon and nitrogen in adult workers using stable isotope analysis. Carbon in adult workers was largely derived from plant sources. Worker nitrogen isotopic ratios varied significantly among colonies and were enriched compared to prey. Prey nitrogen isotope ratios suggest considerable intercolonial variation in diet, with some colonies collecting prey from lower trophic levels than other colonies. The importance of nectar as a source of metabolic carbon in adult worker biomass, coupled with the high frequency of nectar collection, supports the conclusion that omnivory is a key to supporting this species’ biomass in Neotropical wet forests.     

Small-scale variability in a mosaic tropical rainforest influences habitat use of long-tailed macaques

Pristine habitats have generally been considered to be the most important ecological resource for wildlife conservation, but due to forest degradation caused by human activities, mosaics of secondary forests have become increasingly prominent. We studied three forest types in a mosaic tropical forest consisting of short secondary forest (SS), tall secondary forest (TS) and freshwater swamp forest (SF). These forests differed in stand structure and floristic composition, as well as phenological productivity of fruits, flowers and young leaves. We examined habitat use of long-tailed macaques (Macaca fascicularis) in relation to indices of phenological activity. The macaques used the SS for feeding/foraging more than the TS and the SF. This was because the SS had higher productivity of fruit, which is a preferred food resource for macaques. Stem densities of young leaves in the SS and the TS also influenced habitat use, as they provided more clumped resources. Use of SF was limited, but these forests provided more species-rich resources. Our results showed that M. fascicularis responded to small-scale variability in phenological activity between forest types found in a heterogeneous mosaic forest, with young secondary regrowth forests likely providing the most important food resources. Mosaic landscapes may be important as they can buffer the effects of temporal food resource variability in any given forest type. In our increasingly human-altered landscapes, a better understanding of the role of secondary forest mosaics is crucial to the conservation and management of wildlife habitats and the animals they support.     

Species-Specific Effects of Ant Inhabitants on Bromeliad Nutrition

Predator activities may lead to the accumulation of nutrients in specific areas of terrestrial habitats where they dispose of prey carcasses. In their feeding sites, predators may increase nutrient availability in the soil and favor plant nutrition and growth. However, the translocation of nutrients from one habitat to another may depend on predator identity and diet, as well as on the amount of prey intake. Here we used isotopic (¹⁵N) and physiological methods in greenhouse experiments to evaluate the effects of the identity of predatory ants (i.e., the consumption of prey and nest sites) on the nutrition and growth of the bromeliad Quesnelia arvensis. We showed that predatory ants with protein-based nutrition (i.e., Odontomachus hastatus, Gnamptogenys moelleri) improved the performance of their host bromeliads (i.e., increased foliar N, production of soluble proteins and growth). On the other hand, the contribution of Camponotus crassus for the nutritional status of bromeliads did not differ from bromeliads without ants, possibly because this ant does not have arthropod prey as a preferred food source. Our results show, for the first time, that predatory ants can translocate nutrients from one habitat to another within forests, accumulating nutrients in their feeding sites that become available to bromeliads. Additionally, we highlight that ant contribution to plant nutrition may depend on predator identity and its dietary requirements. Nest debris may be especially important for epiphytic and terrestrial bromeliads in nutrient-poor environments.     

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).     

Temporal polyethism in Korean yellowjacket foragers,Vespula koreensis (Hymenoptera,Vespidae)

We examined the foraging behavior of the Korean yellowjacket, Vespula koreensis, to determine whether this species displays temporal polyethism. Using video-recordings of the entrances of artificial nest boxes installed in the field, we investigated the association between the tasks performed by workers and age. We identified three foraging tasks (pulp, nectar and prey foraging). Pulp foraging was performed by younger foragers, while nectar and prey foraging were performed by older foragers. We measured the foraging time (time spent outside of the nest during a single foraging bout) and the weight of the materials that foragers brought into the nest for each task to estimate the cost of the task. Pulp foraging was less costly than nectar or prey foraging by both measures. Taken together, the results suggest that yellowjacket foragers tend to perform low-cost task in their early foraging days and high-cost task later. Our results add to a growing literature showing temporal polyethism in social insects.     

The significance of visual landmarks for navigation of the giant tropical ant,Paraponera clavata (Formicidae,Ponerinae)

Summary Workers of the giant tropical ant,Paraponera clavata, use trail pheromones for orientation and recruitment of nestmates. However, chemical markings may not always be sufficient for successful navigation in complex three-dimensional terrain, and additional orientation cues may be required. Behavioral field experiments were performed to investigate the significance of visual landmarks for homing foragers. Animals which were prevented from seeing the canopy were unable to navigate back to the nest, even though trail pheromones were still present. In contrast, foragers found their way back to the nest after their trail pheromones had been abolished but their visual scenes remained unchanged. This emphasizes the important role of visual landmarks during spatial orientation in homingP. clavata foragers. Individually foraging scouts were discovered in the understory of the forest floor up to 30 m away from their nest. They were rewarded, and displaced between 0.8 m and 13.6 m. Fifteen out of 16 animals had no difficulties in finding the nest entrance despite the altered appearance of local and distant landmarks at the release site. Apparently the scouts were able to recognize the visual scenes at the release site, and used them for reference to locate the nest entrance. In contrast, ants displaced from their nest to sites around 4 m away had more difficulties to re-find the nest.     

Habitat Use and Selection by California Spotted Owls in a Postfire Landscape

ABSTRACT Forest fire is often considered a primary threat to California spotted owls (Strix occidentalis occidentalis) because fire has the potential to rapidly alter owl habitat. We examined effects of fire on 7 radiomarked California spotted owls from 4 territories by quantifying use of habitat for nesting, roosting, and foraging according to severity of burn in and near a 610-km²fire in the southern Sierra Nevada, California, USA, 4 years after fire. Three nests were located in mixed-conifer forests, 2 in areas of moderate-severity burn, and one in an area of low-severity burn, and one nest was located in an unburned area of mixed-conifer-hardwood forest. For roosting during the breeding season, spotted owls selected low-severity burned forest and avoided moderate- and high-severity burned areas; unburned forest was used in proportion with availability. Within 1 km of the center of their foraging areas, spotted owls selected all severities of burned forest and avoided unburned forest. Beyond 1.5 km, there were no discernable differences in use patterns among burn severities. Most owls foraged in high-severity burned forest more than in all other burn categories; high-severity burned forests had greater basal area of snags and higher shrub and herbaceous cover, parameters thought to be associated with increased abundance or accessibility of prey. We recommend that burned forests within 1.5 km of nests or roosts of California spotted owls not be salvage-logged until long-term effects of fire on spotted owls and their prey are understood more fully.     

Foraging strategy predicts foraging economy in a facultative secondary nectar robber

In mutualistic interactions, the decision whether to cooperate or cheat depends on the relative costs and benefits of each strategy. In pollination mutualisms, secondary nectar robbing is a facultative behavior employed by a diverse array of nectar‐feeding organisms, and is thought to be a form of cheating. Primary robbers create holes in floral tissue through which they feed on nectar, whereas secondary robbers, which often lack chewing mouthparts, feed on nectar through existing holes. Because primary robbers make nectar more readily available to secondary robbers, primary robbers facilitate the behaviors of secondary robbers. However, the net effect of facilitation on secondary robber fitness has not been empirically tested: it is unknown whether the benefit secondary robbers receive is strong enough to overcome the cost of competing with primary robbers for a shared resource. We conducted foraging experiments using the bumble bee Bombus bifarius, which can alternatively forage ‘legitimately’ (from the floral opening) or secondary‐rob. We measured the relative foraging efficiencies (handling time per flower, flowers visited per minute, proportion of foraging bout spent consuming nectar) of these alternative behaviors, and tested whether the frequency of primary robbing and nectar standing crop in primary‐robbed flowers of Linaria vulgaris (Plantaginaceae) affected foraging efficiency. Surprisingly, there was no effect of primary robbing frequency on the foraging efficiency of secondary‐robbing B. bifarius. Instead, foraging strategy was a major predictor of foraging efficiency, with legitimate foraging being significantly more efficient than secondary robbing. Legitimate foraging was the more common strategy used by B. bifarius in our study; however, it is rarely used by B. bifarius foraging on L. vulgaris in nature, despite indications that it is more efficient. Our results suggest the need for deeper investigations into why bees adopt secondary robbing as a foraging strategy, specifically, the environmental contexts that promote the behavior.     

Experimental suppression of ants foraging on rainforest vegetation in New Guinea: testing methods for a whole‐forest manipulation of insect communities

1. Ants are extremely abundant in lowland tropical forests where they are important predators, plant mutualists, and herbivores. Their complex role in tropical plant–insect food webs can be best assessed by experimental manipulation of their abundance. Historically, ant exclusion experiments have had a small‐scale focus, such as single trees. Here, we test a new ‘whole‐forest' method of ant exclusion, using treated canopy bait stations, in a diverse tropical rainforest in New Guinea. 2. We conducted a 10‐month manipulative experiment in primary and secondary rainforests. In each forest type, a 625 m² treatment plot was isolated from the surrounding forest and 135 bait stations treated with fipronil, S‐methoprene, and hydramethylnon were placed in trees to suppress ants. Ant activity was monitored in the forest canopy and understorey with an additional 65 stations in treatment and control plots. 3. We achieved a dramatic decline in ant abundance in treatment plots compared with controls in both forest types, with an average decrease in ant numbers per station of 82.4% in primary and 91.2% in secondary forest. In particular, native dominant species Oecophylla smaragdina, Anonychomyrma cf. scrutator in primary forest, and invasive Anoplolepis gracilipes in secondary forest were greatly affected. In contrast, Tapinoma melanocephalum flourished in treatment plots, perhaps benefiting from reduced competition from other ant species. 4. Our study demonstrates that it is possible to selectively eradicate most of the foraging ants in a structurally complex tropical forest. We propose whole‐forest manipulation as a novel tool for studying the role of ants in shaping plant–insect food webs.     

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.     

Ambush Predation of Stingless Bees (<Emphasis Type="Italic">Tetragonisca angustula</Emphasis>) by the Solitary-Foraging Ant <Emphasis Type="Italic">Ectatomma tuberculatum</Emphasis>

Social insect colonies are high-value foraging targets for insectivores, prompting the evolution of complex colony defensive adaptations as well as specialized foraging tactics in social insect predators. Predatory ants that forage on other social insects employ a diverse range of behaviors targeted at specific prey species. Here, we describe a solitary foraging strategy of the ant Ectatomma tuberculatum, on nest guards of the stingless bee Tetragonisca angustula. We observed multiple instances of E. tuberculatum ambushing and successfully capturing the hovering and standing guards of T. angustula near nest entrances. The unique hovering behavior of the guard caste of this bee species, an adaptation to frequent cleptoparasitism by other stingless bees, may make these guards particularly vulnerable to ground-based, ambush attacks by E. tuberculatum. Likewise, the behavior of the foraging ants appears to adaptively exploit the defensive formations and activity patterns of these bees. These observations suggest an adaptive and targeted predatory strategy aimed at gathering external guard bees as prey from these heavily fortified nests.     

Prey‐driven behavioral habitat use in a low‐energy ambush predator

Food acquisition is an important modulator of animal behavior and habitat selection that can affect fitness. Optimal foraging theory predicts that predators should select habitat patches to maximize their foraging success and net energy gain, likely achieved by targeting areas with high prey availability. However, it is debated whether prey availability drives fine‐scale habitat selection for predators. We assessed whether an ambush predator, the timber rattlesnake (Crotalus horridus), exhibits optimal foraging site selection based on the spatial distribution and availability of prey. We used passive infrared camera trap detections of potential small mammal prey (Peromyscus spp., Tamias striatus, and Sciurus spp.) to generate variables of prey availability across the study area and used whether a snake was observed in a foraging location or not to model optimal foraging in timber rattlesnakes. Our models of small mammal spatial distributions broadly predicted that prey availability was greatest in mature deciduous forests, but T. striatus and Sciurus spp. exhibited greater spatial heterogeneity compared with Peromyscus spp. We found the spatial distribution of cumulative small mammal encounters (i.e., overall prey availability), rather than the distribution of any one species, to be highly predictive of snake foraging. Timber rattlesnakes appear to forage where the probability of encountering prey is greatest. Our study provides evidence for fine‐scale optimal foraging in a low‐energy, ambush predator and offers new insights into drivers of snake foraging and habitat selection.     

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

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

Division of labor in post-emergence colonies of the primitively eusocial waspPolistes instabilis de Saussure (Hymenoptera: Vespidae)

Polyethism was quantified in post-emergence colonies of the primitively eusocial wasp,Polistes instabilis, and compared to polyethism in a sympatric advanced eusocial wasp,Polybia occidentalis. Like P.occidentalis, P. instabilis foragers collected food (nectar and prey) and nest materials (wood pulp and water).P. instabilis foragers showed some evidence of specialization with respect to which materials they gathered, but most foragers, divided their effort among food and nest materials, a pattern that is rarely seen inP. occidentalis. In colonies of both species, more foragers collected nectar than any other material; in contrast, most water foraging was performed by one or two workers. Upon returning to the nest,P. instabilis foragers gave up part or all of most nectar, prey, and pulp loads to nestmates, while water was rarely partitioned. Prey loads were most likely to be given up entirely.P. instabilis workers show evidence of conflict over the handling of materials at the nest. The frequency with which workers took portions of nectar loads from forgers was positively correlated with their frequency of aggressive dominant behavior, and with their frequency of taking other foraged materials. Compared to polyethism inP. occidentalis P. instabilis showed less individual specialization on foraging tasks and less partitioning of foraged materials with nestmates, suggesting that these characteristics of polyethism have been modified during the evolution of advanced insect societies.