How do ants assess food volume?

By comparing the behaviour of Lasius niger scouts at sucrose droplets of different volumes, we empirically identified the criterion used by each scout to assess the amount of food available as well as the rules governing its decision to lay a recruitment trail. When scouts discovered food volumes exceeding the capacity of their crop (3 or 6 μl), 90% immediately returned to the nest laying a recruitment trail. In contrast, when smaller food droplets (0.3, 0.7 or 1 μl) were offered, several scouts stayed on the foraging area, presumably exploring it for additional food. If unsuccessful, they returned to the nest without laying a trail. The droplet volume determined the percentage of trail-laying ants but had no influence on the intensity of marking when this was initiated. The key criterion that regulated the recruiting behaviour of scouts was their ability to ingest their own desired volume. This volume acted as a threshold triggering the trail-laying response of foragers. Collective regulation of foraging according to food size resulted from the interplay between the distribution of these desired volume thresholds among colony members and the food volume available. We relate some aspects of the foraging ecology of aphid-tending ants to this decision-making process. Copyright 2000 The Association for the Study of Animal Behaviour.     

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.     

Out of sight but not out of mind: modulation of recruitment according to home range marking in ants

Animals can acquire a global knowledge about their environment that exceeds their individual capacities by estimating the local density and activity of nestmates in an area. In ants, home range marking can indicate the density and activity of nestmates, allowing scouts to assess the potential interest of the area as a foraging site. We investigated how home range marking through footprints influences the foraging behaviour of Lasius niger scouts at a sugary food source (1 M, 1.5 ml). Over a marked apparatus the discovery time of food sources decreased while the probability of scouts recruiting nestmates and of continuing to lay a trail increased. For ants making U turns on their return to the nest, home range marking helped them to resume laying a trail after the U turn and delayed the occurrence of the U turn. As a result, the trail intensity and the rate at which information about food was conveyed by scouts to nestmates depended on home range marking. Such modulation of information reduces the number of foragers mobilized to less frequented areas that are potentially dangerous and promotes recruitment and exploitation of food sources to better known sites.     

Self-organized asymmetries in ant foraging: a functional response to food type and colony needs

The dominant paradigm to explain asymmetries in the spatialdistribution of foraging animals is that they track the spatialheterogeneity of their environment. However, in social insects,endogenous spatial asymmetries can emerge within a uniformenvironment as an outcome from the self-organizing processof trail recruitment. We studied how self-organized asymmetries contribute to the exploitation of different food sources (carbohydrateor proteins) in colonies of the aphid-tending ant Lasius nigervarying in their nutritional needs (presence or absence ofbrood). Colonies with brood fed on sucrose sources exhibita higher mobilization of foragers than the other experimentalgroups. Foraging patterns differ greatly according to food type: colonies strongly focus their activity on only one dropletof sucrose, whereas they show a rather homogeneous distributionof foragers between proteinaceous sources. In addition, thepresence of brood in the colony enhances the asymmetry of collectiveforaging for both types of food. These spatial differencesin self-organized foraging patterns allow efficient exploitationof natural resources and play a role in the competitive strategy of this widespread palearctic ant.     

Regulated food recruitment through individual behavior of scouts in the ant,Myrmica sabuleti (Hymenoptera: Formicidae)

By presenting different kinds of food sources to colonies ofM. sabuleti, we have demonstrated that this species regulates its foraging activity by altering the proportion of scouts that return to the nest to recruit nestmates after discovering a food source and by varying the number of nestmates recruited by a scout. These two parameters are related to the kind of food discovered. Our behavioral experiments showed that the probability that a scout returned to the nest decreased with a decreasing quantity of sucrose solution. In contrast, the number of returned scouts that elicited recruitment from the nest and the mean number of nestmates recruited by one of these scouts were independent of the quantity of the sucrose solution. Recruitment even occurred toward a 1- or 0.25-µl droplet of sucrose solution. When a scout discovered a large dead prey, a large drop of prey juice, a cluster of 30 dead fruit flies, or 1 isolated fruit fly, it always went back to the nest, but it elicited recruitment only when the food source was a large dead prey or a large drop of prey juice. No recruitment occurred when the food source was a single fruit fly and recruitment occurred only once in 30 trials when a cluster of 30 fruit flies was discovered.     

Individual flexibility and choice of foraging strategy in Polyrhachis laboriosa F. Smith (Hymenoptera, Formicidae)

Summary: We report in this study that the tree-dwelling African ant Polyrhachis laboriosa (Formicinae) uses different foraging strategies according to the size of the available food sources. We demonstrate that a recruitment behaviour can be induced with a 125 7l alimentary reward and that foraging remains solitary when rewards are smaller. Small rewards do not elicit trail-laying behaviour, and exploration behaviour is considerable. With large permanent food sources, scouts use group recruitment and there is less exploration around the reward. The choice of the foraging strategy is determined by the first forager, which modifies its behaviour according to the volume of the food supply. Independently of the size of the reward, the forager shows many exploratory displays during the first visit to the source, and contrary to most ants, it never lays a trail during its first return to the nest. Visual cues remain mainly used for individual orientation; information collected during the first trips are then transmitted to nestmates thanks to temporary trail laying behaviour.     

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.     

How patrollers set foraging direction in harvester ants

Recruitment to food or nest sites is well known in ants; the recruiting ants lay a chemical trail that other ants follow to the target site, or they walk with other ants to the target site. Here we report that a different process determines foraging direction in the harvester ant Pogonomyrmex barbatus. Each day, the colony chooses from among up to eight distinct foraging trails; colonies use different trails on different days. Here we show that the patrollers regulate the direction taken by foragers each day by depositing Dufour's secretions onto a sector of the nest mound about 20 cm long and leading to the beginning of a foraging trail. The patrollers do not recruit foragers all the way to food sources, which may be up to 20 m away. Fewer foragers traveled along a trail if patrollers had no access to the sector of the nest mound leading to that trail. Adding Dufour's gland extract to patroller-free sectors of the nest mound rescued foraging in that direction, while poison gland extract did not. We also found that in the absence of patrollers, most foragers used the direction they had used on the previous day. Thus, the colony's 30-50 patrollers act as gatekeepers for thousands of foragers and choose a foraging direction, but they do not recruit and lead foragers all the way to a food source.     

Trail Communication During Foraging and Recruitment in the Subterranean Termite Reticulitermes santonensis De Feytaud (Isoptera, Rhinotermitidae)

The search for food in the French subterranean termite Reticulitermes santonensis De Feytaud is organized in part by chemical trails laid with the secretion of their abdominal sternal gland. Trail-laying and -following behavior of R. santonensis was investigated in bioassays. During foraging for food termites walk slowly (on average, 2.3 mm/s) and lay a dotted trail by dabbing the abdomen at intervals on the ground. When food is discovered they return at a quick pace (on average, 8.9 mm/s) to the nest, laying a trail for recruiting nestmates to the food source. While laying this recruitment trail the workers drag the abdomen continuously on the ground. The recruitment trail is highly attractive: it is followed within a few seconds, by more nestmates, and at a quicker pace (on average, 6.4 mm/s) than foraging trails (on average, 2.9 mm/s). The difference between foraging and recruitment trails in R. santonensis could be attributed to different quantities of trail pheromone. A caste-specific difference in trail pheromone thresholds, with workers of R. santonensis being more sensitive to trails than soldiers, was also documented: soldiers respond only to trails with a high concentration of trail pheromone.     

Influence of seed size and seed nature on recruitment in the polymorphic harvester ant Messor barbarus

In the polymorphic harvester ant Messor barbarus, the colony modulates response during foraging according to seed characteristics. The easiness of picking up small seeds (oat fragments or canaryseeds) shortens the time lapse between food discovery and the return of scouts to the nest, favouring the onset of recruitment and higher mobilisation rates than bigger seeds (whole oat). However, size is not the only criterion, as canaryseeds are preferred among small ones. A modulation of the trail laid by the first scouts also plays a role in the shaping of harvesting patterns and in the enhancement of recruitment towards small and/or preferred seed. This flexibility extends to the relative participation of the three worker sizes classes, which differs according to seed. Media are the most numerous at the foraging arena and largely responsible for trail-laying. Minor participate in seed harvesting and trail-laying, mostly when they can be efficient in carrying seeds, such as oat fragments. Major scarcely participate in harvesting and trail-laying, but are involved in the exploitation of bigger and/or preferred seed species. When faced with different seed species, M. barbarus seems thus to collectively adopt a “foraging time minimisation” rather than an “energetic gain per item harvested maximisation” strategy. Dynamics of foraging and division of labour between foragers of different sizes are related to both the workers’ ability to carry seed items, and the transfer of information to nestmates by trail-laying, all parameters that vary according to seed characteristics.     

Tandem Recruitment and Trail Laying in the Ponerine Ant Diacamma rugosum: Signal Analysis1

During nest emigration, Diacamma rugosum scouts recruit female nestmates to the new nest site by tandem running. ♂♂, brood and prey objects are carried. The tandem leaders stimulate their nestmates to follow them by pushing or, more rarely, pulling them with the mandibles. The signals releasing the following behaviour are mainly chemical. Only 15–30% of motivated followers react positively to an odourless paper or wax dummy. The ♀♀ follow cut-off thoraces, legs and gasters plugged at the end or without hind gut and sting apparatus, just as well as a leading ant. The ants even follow a dummy consisting of body parts if they are not in direct mechanical contact with it. The leading effect of the body surface is not reduced by intensive cleaning with water or by extraction in solvents of different polarity. Only maceration in potash lye eliminates the leading effectiveness of the cuticula. None of the secretions of the tested abdominal glands release following behavior. Only the hind gut fluid possesses leading signal qualities. No distribution of faeces over the bodies of the leading ants can be observed before or during tandem running. The faecal pheromone responsible for this leading effect does not seep into the superficial layer of a leader dummy made from bee wax but it can be washed off with water. These results show that the actual signal releasing tandem following is an unspecific odour substance of the cuticula surface. The presence of the hind gut fluid is not necessary for the tandem following response. It may be an additional signal. During nest emigration the hind gut fluid is scattered patchily in the form of small droplets by the nest finders on their way to the new nest. No increased deposition of faecal droplets could be observed on the way to repeatedly visited rich food sources. The hind gut fluid serves as a long lasting trail substance and has an exclusively orienting effect. The secretions of the other abdominal glands are not trail pheromones. The nest entrance is marked colony-specifically.     

Food availability affects<Emphasis Type="Italic"> Osmia pumila</Emphasis> (Hymenoptera: Megachilidae) foraging,reproduction, and brood parasitism

Food limitation can reduce reproductive success directly, as well as indirectly, if foraging imposes a risk of predation or parasitism. The solitary bee Osmia pumila suffers brood parasitism by the cleptoparasitic wasp Sapyga centrata, which enters the host nest to oviposit while the female bee is away. I studied foraging and reproduction of O. pumila nesting within cages stocked with rich or sparse floral resources, and the presence or absence of S. centrata to test (1) the response of nesting female O. pumila to food shortages, (2) the response of nesting female O. pumila to the presence of parasites, and (3) whether brood produced under scarce resources are more likely to be parasitized by S. centrata. The rate of brood cell production was significantly lower in cages with sparse floral resources, although females in sparse cages did not produce significantly fewer brood cells overall. Sapyga centrata did not influence the rate of brood cell production, but females exposed to the cleptoparasites had marginally significantly lower reproductive output. Nests in parasite cages had significantly fewer brood cells than those in parasite free cages. The mean duration of foraging bouts made by female O. pumila in sparse cages was not significantly longer than that in rich cages. O. pumila spent less time in the nest between pollen and nectar foraging bouts in sparse cages with S. centrata than those in other cages suggesting that these individuals made more frequent food foraging trips. Despite the weak effects of parasites and bloom density on foraging behavior, O. pumila brood cells experienced a 5-fold higher probability of parasitism by S. centrata in cages with sparse bloom than in those with rich bloom [corrected]. These results support the hypothesis that indirect effects of food scarcity increase O. pumila susceptibility to brood parasitism, although the exact mechanism is not entirely clear yet.     

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.     

Flexibility of Individual Load-mass Selection in Relation to Foraging Trail Gradient in the Leaf-cutter Ant Acromyrmex octospinosus

The stochasticity in food quality and availability, and physical trail characteristics experienced by leaf-cutter ants, may favour individual flexibility in load-mass selection so as to forage effectively. The present study aimed to confirm previous evidence, from Atta cephaoltes foragers, of variable load-mass selection in response to steep inclines and declines in the leaf-cutter ant Acromyrmex octospinosus. The foraging trail gradient of a captive colony of Ac. octospinosus was manipulated by altering the position of a foraging platform relative to the nest box. The results indicate an effect of steep gradients on walking speed and variation in load mass in relation to gradient as a result of individual plasticity, not recruitment of different-sized individuals. Ants selected heavier loads when returning to the nest vertically downwards than when returning horizontally or vertically upwards. These results are discussed with reference to foraging performance. Walking speed was considerably reduced on upward returns to the nest, but was also slower when travelling vertically downwards compared with horizontal trails, suggesting vertical trails per se impact on the time costs of foraging. Differences in load-mass selection were evident from the onset of foraging and did not change significantly over the course of 24 h, suggesting this behaviour was based on individual experience, rather than colony-level information feedback. The present study has demonstrated that Ac. octospinosus foragers are capable of individual flexibility in load-mass selection in response to a physical trail characteristic that is pertinent to their natural habitat and is a factor seldom considered in theoretical foraging models.     

Group hunting in a ponerine ant,Leptogenys nitida Smith

Field observations on the emigration and foraging behaviours of the southern African ponerine ant, Leptogenys nitida, were undertaken at Mtunzini, Natal, South Africa. These colonies have a single ergatoid queen and 200–1000 workers. The nest sites are found in the leaf litter and these nests are moved frequently over distances ranging from 0.5 to 5 m. Leptogenys nitida is a diurnal predator of arthropods dwelling in the leaf litter. Up to 500 workers participate in each foraging trail, and are not led by definite scouts. Ants form clear trunk trails and fan out at various intervals to search for prey. The prey is searched for and retrieved cooperatively. From laboratory tests it was determined that ants will follow pygidial gland extracts, with the poison gland extract eliciting a limited response. The type of army ant behaviour observed in L. nitida seems to be different to that observed in other ponerine ants.     

Nest demographics and foraging behavior of <Emphasis Type="Italic">Apterostigma collare</Emphasis> Emery (Hymenoptera,Formicidae) provide evidence of colony independence

Apterostigma collare Emery is a highly derived fungus-growing ant within the Tribe Attini whose small, fungal nests are found in tropical rain forests. This study focuses on determining the colony structure of A. collare, specifically searching for evidence of polydomy or independence. We surveyed and observed nests in the field, and performed foraging bioassays and dissected nests in the laboratory. We determined the size and contents of nests in field populations. Nests found near other nests were not statistically different in size compared to nests found alone. There was also no statistical difference between near and lone nests regarding the presence of a queen in the nest. Most nests contained one queen with brood and workers, regardless of their proximity to other nests. Observations also were made of foraging and trail-marking behaviors. Foraging activity observed in the field revealed that workers left the nest area and followed trails upwards into the canopy, but they did not interact with foragers from other nearby nests. In a laboratory foraging arena, foragers marked a trail to a food source by dragging the gaster. Bioassays showed that A. collare workers preferred their own foraging trails, but not those of other conspecific colonies. All results suggest that each nest represents an independent colony, supporting a previous report that nests found in close proximity do not constitute a polydomous colony. Received 19 July 2006; revised 23 March 2007; accepted 6 June 2007.     

The role of pheromones in the initiation of foraging, recruitment and defence by the soldiers of a tropical termite, Nasutitermes corniger (Motschulsky)

The polyethic behaviour of soldiers and workers of Nasutitermescorniger during foraging and in defence is described. It hasbeen shown that the cephalic gland defensive secretion of thenasute soldiers produces short term recruitment of only soldiersto loci where it is present. Without further reinforcement ofthe stimulus, the number of soldiers wanes. Workers react tothe cephalic gland secretion by remaining in or retreating tothe nest. The sternal gland secretion of workers and soldiersacts in long term recruitment of soldiers when presented experimentallyas a point source, as a trail or as a trail with a food sourceat its end. This secretion causes more exiting soldiers thanthe cephalic gland secretion, and is more effective in thisregard as a trail than as a point source. The greater numbersexiting in the trail situation appears to be related to thegreater amount of pheromone area present, and the halting ofsoldiers at the pheromone odour boundary. The greatest numberof soldiers exit when the trail is coupled with a food sourceat its end, and in this situation workers exit for the firsttime in significant numbers. Additional information about thefood source is postulated to be communicated by tactile means(jittering) and it is suggested that this lowers the thresholdfor the trail following responses in the termites such thatworker exiting is initiated. Other factors affecting the numbersforaging are noted. The phenomenon of soldier initiation offoraging as scouts, and as a scouting column, is commented onwith regard to the polyethic responses to the pheromones, andto its adaptive significance (sociobiological import) in thelight of the soldiers' rôle in colony defence.     

Trail laying behaviour during food recruitment in the antLasius niger (L.)

Summary The trail-laying behaviour of foragers of the antLasius niger was observed in the laboratory on a 20 cm bridge between the nest and the food source. We measured both the frequency of trail laying, as defined by the proportion of trips during which trail laying occurred, and its intensity, as defined by the number of marks laid during one bridge crossing.Foragers do not exhibit trail-laying behaviour until a food source is discovered. Trail laying then occurs more or less equally both to and from the nest, and both its frequency and intensity decrease as the recruitment proceeds. Foragers from very small colonies less than a year old appear to have quantitatively the same trail laying behaviour as those from older and much larger colonies.Groups of recruiters and recruits were individually marked. Their trail laying intensity was similar, both for trips to and from the nest, and for an ant's first, second, third and fourth trip. The frequency diminished rapidly with the number of trips made by each individual, and was 2–3 times higher for recruiters than for recruits, for trips both to and from the nest. Even though foragers stop marking after a variable number of passages, they continue to move between the nest and the food source, and other ants start marking. Different foragers appear to have widely different levels of trail laying, although we cannot say whether these differences are stable between different recruitments.Trail laying is strongly affected by the foragers' position on the bridge, especially for ants returning to the nest which lay up to five times more on the segment closest to the source than that closest to the nest. Foragers on a weakly marked trail appear to mark more than those on a well-marked trail. However, this effect is weak and could partly be attributed to their lower speed.Finally, a model using the experimental data gathered on the individuals' trail-laying behaviour reproduced satisfactorily the colony's overall trail laying.     

Ants use pheromone markings in emigrations to move closer to food-rich areas

Nest site quality can affect survival and reproduction, and thus many animals have evolved behaviors which facilitate nest site assessment and selection. Ants of the genus Temnothorax have been shown to include an array of nest site attributes when choosing such a site. Here, we show that they also include traits of the habitat surrounding nest sites. In particular, we found that during emigration, ants preferred to move to nests located close to a previously explored food-rich area. We also determined that scent markings played a role in this choice and that scouts and transporting ants may have tracked scent marks laid in foraging, and this behavior could have biased emigration toward nests located near previously foraged areas. These results indicate that pheromones play a bigger role in Temnothorax foraging and decision making in emigration than previously thought. Overall, this work provides new insights into the mechanisms involved in habitat selection in ants and contributes to our understanding of collective behavior in social insects in general.     

Comparing Reversal-Learning Abilities,Sucrose Responsiveness,and Foraging Experience Between Scout and Non-Scout Honey bee (Apis mellifera) Foragers

Honey bee (Apis mellifera) colonies divide foraging activities between scouts, who search for new sources of food, and non-scouts, who rely on information from waggle dances to find food sources. Molecular analyses of scouts and non-scouts have revealed differences in the expression of numerous genes, including several related to neurotransmitter signaling. Despite this progress, we know almost nothing about cognitive, sensory, or behavioral differences that underlie scouting. We tested three hypotheses related to differences between scouts and non-scouts. First, we tested whether scouts and non-scouts differ in their reversal-learning abilities and found that scouts showed a significantly faster reversal in their response to an odor that was punished and then rewarded. The results also suggested an interaction between the effects of foraging role (scout or non-scout) and seasonal effects on reversal-learning abilities. Second, we tested whether scouts and non-scouts show differences in responsiveness to sucrose rewards and found no difference. Third, we hypothesized that scouts have more foraging experience than do non-scouts. We tested this by comparing wing damage and found that non-scouts showed greater wing damage in the early summer but not the late summer. Together, these three results contribute to our understanding of the underpinnings of scouting behavior.