How does food distance influence foraging in the ant Lasius niger: the importance of home-range marking

We study the influence of food distance on the individual foraging behaviour of Lasius niger scouts and we investigate which cue they use to assess their distance from the nest and accordingly tune their recruiting behaviour. Globally, the number of U-turns made by scouts increases with distance resulting in longer travel times and duration of the foraging cycle. However, over familiar areas, home-range marking reduces the frequency and thereby the impact of U-turns on foraging times leading to a quicker exploitation of food sources than over unmarked set-ups. Regarding information transfer, the intensity of the recruitment trail reaching the nest decreases with increasing food distance for all set-ups and is even more reduced in the absence of home-range marking. Hence, the probability of a scout continuing to lay a trail changes along the homeward journey but in a different way according to home-range marking. Over unexplored setups, at a given distance from the food source, the percentage of returning trail-laying ants remains unchanged for all tested nest-feeder distances. Hence, the tuning of the trail recruiting signal by scouts was not influenced by an odometric estimate of the distance already travelled by the ants during their outward journey to the food. By contrast, over previously explored set-ups, a distance-related factor – that is the intensity of home-range marking – strongly influences their recruiting behaviour. In fact, over a home-range marked bridge, the probability of returning ants maintaining their trail-laying behaviour increases with decreasing food distance while the gradient of home-range marks even induces ants which have stopped laying a trail to resume this behaviour in the nest vicinity. We suggest that home-range marking laid passively by walking ants is a relevant cue for scouts to indirectly assess distance from the nest but also local activity level or foraging risks in order to adaptively tune trail recruitment and colony foraging dynamics. Received 13 July 2004; revised 26 January and 20 May 2005; accepted 2 July 2005.     

Factors affecting distribution of the mound-building ant Lasius neoniger (Hymenoptera: Formicidae) and implications for management on golf course putting greens

Lasius neoniger (Emery), a cosmopolitan ant species, can be a serious pest when its mound-building activities occur on golf course putting greens and other closely mowed turfgrass sites. We mapped the distribution of 735 ant mounds on 30 sand-based putting greens of three golf courses. We then examined factors that might explain why >90% of the mounds on such greens were concentrated in a 2-m wide band just inside the perimeter. Root aphids (Homoptera: Aphididae) from which L. neoniger obtains honeydew were largely absent from high-sand root zone mix of greens but present in surrounding turfgrass on natural soil. Main ant nests, with brood, also were absent from sand-based greens but abundant in adjacent roughs. Although more root aphids were found within ant nests than away from nests, their numbers seem too low to be the main factor restricting the ants' distribution to edges of putting greens. In manipulative experiments, ants responded to low cut (scalped) turf and to sand-filled holes by increased mound building. We suggest that most ant mounds on sand-based greens are associated with subnests, used by foraging workers, which are connected to main nests located just outside the collar in natural soil. Encroachment of mounds into greens occurs when the polydomous colonies seasonally expand their foraging territories, accounting for mounds being concentrated around the perimeter. Control actions for L. neoniger on golf courses should focus on the perimeter of sand-based greens.     

The foraging ecology of the army ant Eciton rapax: an ergonomic enigma?

Abstract. 1. The army ant Eciton rapax (F. Smith) produces longer raid systems than any other member of its genus and it is a specialist predator of forest floor and understory ants such as species of Camponotus, Odontomachus and Pachycondyla.
2. Allometrical analysis confirms that E. rapax is the only member of its genus without distinct physical castes among its workers: its foraging population is entirely monomorphic and there are no majors.
3. The workers of E. rapax are distributed over a considerable size range, and there are distinct divisions of labour within these colonies: small workers tend to stay in the nests and among the larger foraging workers those retrieving prey items are significantly bigger than the rest.
4. An analysis of foraging efficiency and worker performance in E. rapax suggests that transport costs, resulting from the great distances that workers travel during raids and emigrations, are one of the selection pressures that have favoured the evolution of large monomorphic workers in this species.     

Ontogenetic changes in forager polymorphism and foraging ecology in the leaf-cutting ant Atta cephalotes

In the leaf-cutting ant Atta cephalotes (L.) small colonies produce a relatively narrow size-range of small workers, whereas large colonies produce a much wider size-range of workers. In this study, I compared the foraging of four small A. cephalotes colonies (fewer than 5000 workers) with published data on foraging of large colonies to examine how colony size and worker size-range may be related to foraging ecology in leaf-cutting ants. I found that the foraging ecology of small A. cephalotes colonies is very different from that of large colonies. In small colonies, a relatively narrow size-range of foragers (1.4–6.7 mg, mean 3.3 mg) cut primarily herbs (ferns, grasses, and other small herbaceous plants) located within 7 m of the nest. In contrast, in large colonies, a broader size-range of workers (1.4–30 mg, mean 7.3 mg) participate in foraging, generally harvesting from trees 20–80 m from the nest, with larger workers cutting on trees with thicker and tougher leaves. Small colonies' dependence on small herbaceous plants near the nest may have a profound impact on distribution of A. cephalotes. A. cephalotes colonies are rarely found in primary forest, where the low occurrence of small herbaceous plants in the understory may preclude the establishment of young colonies.     

Spatial patterns of granivorous ant nest abundance and nest site selection in agricultural landscapes of Central Spain

Summary Patterns of abundance and site selection of granivorous ant nests were investigated in extensive cereal croplands of Central Spain. Nest densities and distributions were measured in two consecutive summers (1988 and 1989), together with habitat physiognomy and seed availability. Nest site selection patterns were analysed at two spatial scales (landscape and microhabitat) with respect to habitat physiognomy. Results indicate a very constant and predictable pattern of both nest abundance and nest site selection. Granivorous ant nests were most abundant in shrublands, and shrubby microsites were selected for nest placement. Croplands, and microsites with high covers of bare ground and litter, were avoided. These patterns were consistent between years despite a 1.7-fold increase in shrubland nest densities, that was attributed to the exceptionally dry winter between nest censuses. I suggest that winter survivorship of ant nests in the unploughed landscape units, and periodic ploughing in croplands, may be the main factors constraining granivorous ant densities in the landscape studied.     

Respiration in the sexuals of the ant Lasius flavus

ABSTRACT. Gynes and males of Lasius flavus Fab. were taken from the nest as they eclosed, and at intervals thereafter until the mating flight, and their respiration was measured. After the mating flight respiration was measured in gynes at various stages of colony foundation. Respiration in gynes rose rapidly to a peak as they began to accumulate food reserves for colony foundation. Subsequently, the respiratory rate declined and remained at a low level until the mating flight and during the claustral phase of colony foundation. Once the colony was well established and the queen actively laying, the respiratory rate returned to that at the beginning of maturation. Males showed smaller changes in their respiratory rates, which were generally higher than those of the gynes. The variation in respiration in males seemed to be due to increasing restlessness as the time of the mating flight approached.     

Extended phenotypes and foraging restrictions: ant nest entrances and resource ingress in leaf‐cutting ants

Several factors may restrict the acquisition of food to below the levels predicted by the optimization theory. However, how the design of structures that animals build for foraging restricts the entry of food is less known. Using scaling relationships, we determined whether the design of the entrances of leaf‐cutting ant nests restricts resource input into the colony. We measured nests and foraging parameters in 25 nests of Atta cephalotes in a tropical rain forest. Ant flux was reduced to up to 60% at nest entrances. The width of all entrances per nest increased at similar rates as nest size, but the width of nest entrances increased with the width of its associated trail at rates below those expected by isometry. The fact that entrance widths grow slower than trail widths suggests that the enlargement of entrance holes does not reach the dimensions needed to avoid delays when foraging rates are high and loads are big. The enlargement of nest entrances appears to be restricted by the digging effort required to enlarge nest tunnels and by increments in the risk of inundation, predator/parasitoid attacks and microclimate imbalances inside the nest. The design of the extended phenotypes can also restrict the ingress of food into the organisms, offering additional evidence to better understand eventual controversies between empirical data and the foraging theory. Abstract in Spanish is available with online material.     

Unpaved roads alter foraging patterns of the leafcutter ant Atta colombica

Leaf cutting ants are dominant herbivores and influential ecosystem engineers in the Neotropics. It has been suggested that habitat disturbances alter the architecture of foraging trail systems for colonies in their vicinity; however, the evidence remains scarce. In this study we investigated the effect of unpaved roads dissecting tropical lowland forest habitat on the structure of leafcutter foraging trail systems and foraging effort. We mapped trail systems for 16 mature Atta colombica colonies located at different distances from unpaved roads. Our results suggest exploitation of unpaved roads by leafcutters provides favorable foraging conditions, causing significant differences in foraging trail structure.     

Collective exploration and area marking in the ant Lasius niger

Summary: The aphid tending ant species, Lasius niger collectively explore new areas. Scouts are mobilised within the first five minutes of exploration with activity thereafter decreasing and stabilising after 40 minutes. Collective path choices of colonies underline the existence of chemical recruitment and area marking during exploration. To identify the territorial or home-range role of exploratory marking, we examined its influence on agonistic behaviour during intraspecific encounters. Ten nestmates and ten alien conspecifics were confronted: (i) on a paper marked by their own colony, (ii) on a paper marked by the alien colony or (iii) on a paper not chemically marked by ants. Residents and intruders showed a weak level of aggression and behaved similarly whatever the type of area marking. As no colony-specificity was observed in the odds of eliciting aggression, exploratory area marking in Lasius niger seems to be rather a home-range than a true territorial marking. We discuss the role of collective exploration, home-range marking and intraspecific interactions in the structuring and use of foraging space by neighbour Lasius niger colonies.     

Queen mating and paternity variation in the ant Lasius niger

We have electrophoretically analysed the variation in queen mating and worker paternity across and within seven populations of the ant Lasius niger in northwestern Europe. Populations were panmictic and not genetically differentiated ( F _ST = 0.003 ± 0.004; range c. 1000 km). Queens ( n = 535) were shown to mostly mate with a single male, but double mating occurred in all populations and triple mating was found in one case (the total number of worker offspring analysed for paternity was 4825). The genetically effective queen mating frequency was 1.16 on average across populations (range 1.04–1.42). Double sampling of six out of the seven populations showed that most of the variation in queen mating occurred among populations and not within populations among years. Also, paternity skew in colonies with double-mated queens was relatively constant per site but varied across populations. Paternity skew was high in populations with low frequencies of double queen-mating, low in populations with intermediate frequencies of double queen mating and ambiguous in a single population where more than half of the queens mated multiply. Double-mated queens were only collected halfway through a nuptial flight, suggesting that double mating is time consuming and that the mating swarm sex ratio may affect the likelihood of multiple mating towards the end of a flight. No difference in fresh weight between single- and double-mated queens from the same population was found.     

Diel and seasonal patterns of foraging activity in the arboreal ant Crematogaster matsumurai Forel

The foraging habits of the arboreal ant Crematogaster matsumurai Forel were studied in natural and captive colonies in southern Kyushu, Japan. The nests of C. matsumurai are made mainly in the trunks of living trees. In the natural environment, the species mainly lives in decayed parts of relatively tall trees, such as Acer palmatum, Prunus jamasakura and Prunus yedoensis. Observation over a 24 h period showed that C. matsumurai foraged actively at night as well as during the day. The foraging of the workers out of the nest was seen from early spring to late autumn, both on the tree and on the ground. As protein sources, aphids were mainly collected by foragers from the tree, while small parts of dead insects or other arthropods were mainly collected from the ground surface. The foraging activity of workers is assumed to be influenced by the quantity and quality of food rather than seasonality.     

Effect of aromatic cedar mulch on Argentine ant (Hymenoptera: Formicidae) foraging activity and nest establishment

In the laboratory, Argentine ant, Linepithema humile (Mayr), mortality was positively correlated to the length of an aromatic cedar mulch section that had to be crossed before food could be reached. When ants could access food without crossing the mulch, mortality was not correlated to mulch section length. In the field, Argentine ants showed a tendency to avoid aromatic cedar mulch as a nesting substrate. In plant beds alongside buildings the number of ant nests (pockets containing brood) found was not significantly different between aromatic cedar and cypress mulch. However, when pine straw mulch around oak trees was replaced with aromatic cedar or cypress mulch, a similar number of ant nests was found in the cypress mulch as in the original pine straw, whereas numbers in aromatic cedar mulch were significantly lower. Also, fewer ants were trailing on the trees surrounded by aromatic cedar mulch compared with cypress mulch or the original pine straw. The number of ants attracted to apple jelly baits placed alongside the buildings did not differ between mulch types; neither did the number of ant trails crossing the mulch beds around the buildings. We suggest that aromatic cedar mulch may help control Argentine ants and reduce insecticide input when applied in combination with conventional control methods.     

Density-dependent nest predation in waterfowl: the relative importance of nest density versus nest dispersion

When nest predation levels are very high or very low, the absolute range of observable nest success is constrained (a floor/ceiling effect), and it may be more difficult to detect density-dependent nest predation. Density-dependent nest predation may be more detectable in years with moderate predation rates, simply because there can be a greater absolute difference in nest success between sites. To test this, we replicated a predation experiment 10 years after the original study, using both natural and artificial nests, comparing a year when overall rates of nest predation were high (2000) to a year with moderate nest predation (2010). We found no evidence for density-dependent predation on artificial nests in either year, indicating that nest predation is not density-dependent at the spatial scale of our experimental replicates (1-ha patches). Using nearest-neighbor distances as a measure of nest dispersion, we also found little evidence for "dispersion-dependent" predation on artificial nests. However, when we tested for dispersion-dependent predation using natural nests, we found that nest survival increased with shorter nearest-neighbor distances, and that neighboring nests were more likely to share the same nest fate than non-adjacent nests. Thus, at small spatial scales, density-dependence appears to operate in the opposite direction as predicted: closer nearest neighbors are more likely to be successful. We suggest that local nest dispersion, rather than larger-scale measures of nest density per se, may play a more important role in density-dependent nest predation.     

The evolution of queen pheromones in the ant genus Lasius

Queen pheromones are among the most important chemical messages regulating insect societies yet they remain largely undiscovered, hindering research into interesting proximate and ultimate questions. Identifying queen pheromones in multiple species would give new insight into the selective pressures and evolutionary constraints acting on these ubiquitous signals. Here, we present experimental and comparative evidence that 3‐methylalkanes, hydrocarbons present on the queen's cuticle, are a queen pheromone throughout the ant genus Lasius. Interspecific variation in the chemical profile is consistent with 3‐methylalkanes evolving more slowly than other types of hydrocarbons, perhaps due to differential selection or evolutionary constraints. We argue that the sensory ecology of the worker response imposes strong stabilizing selection on queen pheromones relative to other hydrocarbons. 3‐Methylalkanes are also strongly physiologically and genetically coupled with fecundity in at least one Lasius species, which may translate into evolutionary constraints. Our results highlight how honest signalling could minimize evolutionary conflict over reproduction, promoting the evolution and maintenance of eusociality.     

Complex foraging ecology of the red harvester ant and its effect on the soil seed bank

Granivory is an important interaction in the arid and semi-arid zones of the world, since seeds form an abundant and nutritious resource in these areas. While species of the genus Pogonomyrmex have been studied in detail as seed predators, their impact on seed abundance in the soil has not yet been explored in sufficient depth. We studied the impact of the harvesting activities of the ant Pogonomyrmex barbatus on seed abundance in the soil of the Zapotitlán valley, Mexico. We found that P. barbatus activity significantly impacts the abundance of seeds in the soil, which is lower in the sites where P. barbatus forages than it is in sites with no recorded foraging. We also found that P. barbatus distributes intact seeds of three tree species, two of which are nurse plants, and could consequently be promoting the establishment of these species. Using tools derived from graph theory, we observed that the ant-seed interactions exhibit a nested pattern; where more depredated seed species seem to be the more spatially abundant in the environment. This study illustrates the complex foraging ecology of the harvester ant P. barbatus and elucidates its effect on the soil seed bank in a semi-arid environment.     

Triggering and persistence of trail-laying in foragers of the ant Lasius niger

In the ant Lasius niger, the ability to ingest their own desired volume is the key criterion that rules the recruiting behaviour of scouts. This volume acts as a threshold triggering the trail-laying response of foragers. In this paper, we show that this desired volume is specific to each individual and is kept constant over successive trips to a food source. This individual specificity contrasts with the variability of all individual desired volumes within the colony. In this study, it is also shown that, among L. niger foragers, 14% never participate in the formation of the chemical pathway and never lay a trail over successive trips. Among the others foragers, interindividual differences in the persistence of trail-laying behaviour over successive trips are observed but do not rely on an individual specialisation, in which some ants would lay a trail more frequently and persistently than other scouts. We discuss how an individual in the foraging behaviour can play an essential role in the regulation of food retrieval dynamics.     

Hierarchical networks of food exchange in the black garden ant Lasius niger

In most eusocial insects, the division of labor results in relatively few individuals foraging for the entire colony. Thus, the survival of the colony depends on its efficiency in meeting the nutritional needs of all its members. Here, we characterize the network topology of a eusocial insect to understand the role and centrality of each caste in this network during the process of food dissemination. We constructed trophallaxis networks from 34 food-exchange experiments in black garden ants (Lasius niger). We tested the influence of brood and colony size on (i) global indices at the network level (i.e., efficiency, resilience, centralization, and modularity) and (ii) individual values (i.e., degree, strength, betweenness, and the clustering coefficient). Network resilience, the ratio between global efficiency and centralization, was stable with colony size but increased in the presence of broods, presumably in response to the nutritional needs of larvae. Individual metrics highlighted the major role of foragers in food dissemination. In addition, a hierarchical clustering analysis suggested that some domestics acted as intermediaries between foragers and other domestics. Networks appeared to be hierarchical rather than random or centralized exclusively around foragers. Finally, our results suggested that networks emerging from social insect interactions can improve group performance and thus colony fitness.