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.     

Stingless bees (<Emphasis Type="Italic">Scaptotrigona pectoralis</Emphasis>) learn foreign trail pheromones and use them to find food

Foragers of several species of stingless bees (Hymenoptera, Apidae and Meliponini) deposit pheromone marks in the vegetation to guide nestmates to new food sources. These pheromones are produced in the labial glands and are nest and species specific. Thus, an important question is how recruited foragers recognize their nestmates’ pheromone in the field. We tested whether naïve workers learn a specific trail pheromone composition while being recruited by nestmates inside the hive in the species Scaptotrigona pectoralis. We installed artificial scent trails branching off from trails deposited by recruiting foragers and registered whether newly recruited bees follow these trails. The artificial trails were baited with trail pheromones of workers collected from foreign S. pectoralis colonies. When the same foreign trail pheromone was presented inside the experimental hives while recruitment took place a significant higher number of bees followed the artificial trails than in experiments without intranidal presentation. Our results demonstrate that recruits of S. pectoralis can learn the composition of specific trail pheromone bouquets inside the nest and subsequently follow this pheromone in the field. We, therefore, suggest that trail pheromone recognition in S. pectoralis is based on a flexible learning process rather than being a genetically fixed behaviour.     

Homing in Harvester Termites: Evidence of Magnetic Orientation

Colonies of Trinervitermes geminatus (Wasman) build a network of forked foraging trails. Homing workers choose, with few exceptions, those tracks at the forks of the pheromone trail network that lead direct to the nest. Assessments show that these workers are also able to distinguish between the homeward and outward direction along an unbranched trail. Experiments involving the alteration of the local natural magnetic field by means of a weak permanent magnet provide evidence that the termites recognise the home direction by means of magnetoreception.     

Perceptual differences in trail-following leaf-cutting ants relate to body size

Leaf-cutting ants of the genus Atta have highly size-polymorphic workers, and size is related to division of labor. We studied trail-following behavior of different-sized workers in a laboratory colony of Atta vollenweideri. For small and large workers, we measured responsiveness and preference to artificial conspecific and heterospecific pheromone trails made from poison gland extracts of A. vollenweideri and A. sexdens. Responsiveness was measured as the probability of trail-following, and preference was measured by testing the discrimination between one conspecific and one heterospecific trail. Minute amounts of the releaser component methyl-4-methylpyrrole-2-carboxylate (0.4pg/1m), present in both, conspecific and heterospecific trails, suffice to elicit trail-following behavior. Workers followed heterospecific trails, and these trails (after normalizing their concentration) were as effective as conspecific trails. Small workers were less likely to follow a trail of a given concentration than large workers. In the discrimination test, small workers preferred the conspecific trail over the heterospecific trail, whereas large workers showed no significant preference. It is suggested that large workers primarily respond to the releaser component present in both trails, whereas small workers focus more on the conspecific traits provided by the blend of components contained in the trail pheromone.     

Trail‐sharing among tropical ants: interspecific use of trail pheromones?

1. Trail‐sharing between different ant species is rare and restricted to a small number of species pairs. Its underlying mechanisms are largely unknown. For trail‐sharing to occur, two factors are required: (i) one or both species must recognise the other species or its pheromone trails and (ii) both species must tolerate each other to a certain extent to allow joint use of the trail. A species that follows another's trails can efficiently exploit the other's information on food sources contained in the pheromone trails. Hence, food competition and thus aggressive interactions between a species following another's trail and the species being followed, seem likely. 2. In the present study, we investigated interspecific trail following and interspecific aggression in trail sharing associations (i) among Polyrhachis ypsilon, Camponotus saundersi, and Dolichoderus cuspidatus, and (ii) among Camponotus rufifemur and Crematogaster modiglianii. We tested whether trail‐sharing species follow each other's pheromone trails, and whether the ants tolerated or attacked their trail‐sharing partners. In both associations, we confronted workers with pheromone trails of their associated species, and, for the former association, measured interspecific aggression among the trail‐sharing species. 3. In our assays, D. cuspidatus and C. rufifemur regularly followed heterospecific pheromone trails of P. ypsilon and C. modiglianii, respectively. However, only few workers of the remaining species followed heterospecific pheromone trails. Thus, shared trails of P. ypsilon and C. saundersi cannot be explained by interspecific trail‐following. 4. Interspecific aggression among P. ypsilon, C. saundersi, and D. cuspidatus was strongly asymmetric, C. saundersi being submissive to the other two. All three species differentiated between heterospecific workers from the same or another site, suggesting habituation to the respective trail‐sharing partners. We therefore hypothesise that differential tolerance by dominant ant species may be mediated by selective habituation towards submissive species and this way determines the assembly of trail‐sharing associations.     

Trail Pheromone of the Argentine Ant,Linepithema humile (Mayr) (Hymenoptera: Formicidae)

The Argentine ant (Linepithema humile) is recognized as one of the world''s most damaging invasive species. One reason for the ecological dominance of introduced Argentine ant populations is their ability to dominate food and habitat resources through the rapid mobilization and recruitment of thousands of workers. More than 30 years ago, studies showed that (Z)-9-hexadecenal strongly attracted Argentine ant workers in a multi-choice olfactometer, suggesting that (Z)-9-hexadecenal might be the trail pheromone, or a component of a trail pheromone mixture. Since then, numerous studies have considered (Z)-9-hexadecenal as the key component of the Argentine ant trails. Here, we report the first chemical analyses of the trails laid by living Argentine ants and find that (Z)-9-hexadecenal is not present in a detectible quantity. Instead, two iridoids, dolichodial and iridomyrmecin, appear to be the primary chemical constituents of the trails. Laboratory choice tests confirmed that Argentine ants were attracted to artificial trails comprised of these two chemicals significantly more often than control trails. Although (Z)-9-hexadecenal was not detected in natural trails, supplementation of artificial dolichodial+iridomyrmecin trails with an extremely low concentraion of (Z)-9-hexadecenal did increase the efficacy of the trail-following behavior. In stark contrast with previous dogma, our study suggests that dolichodial and iridomyrmecin are major components of the Argentine ant trail pheromone. (Z)-9-hexadecenal may act in an additive manner with these iridoids, but it does not occur in detectable quantities in Argentine ant recruitment trails.     

Fire ant trail pheromones: Analysis of species specificity after gas chromatographic fractionation

The specificity of the trail pheromones of four Solenopsis species was determined using natural trails. Dufour's gland extracts, and purified fractions from Dufour's gland extracts collected after gas-chromatographic separation. S. richteri and S. invicta possess species-specific major trail pheromones, while S. geminata and S. xyloni appear to have a common trail pheromone. Preliminary chemical characterization of the main trail pheromone of S. richteri indicates a M.W. of 218 and empirical formula of C₁₆H₂₆. The trail pheromone system of S. richteri consists of a blend of compounds and this phenomenon may also occur in the other species. The lowest concentration of their trail pheromone that workers of S. richteri could detect was about 10 fg per cm. The significance of blends of pheromones being utilized to generate chemical trails is discussed.     

Chemical and behavioural studies of the trail-following pheromone in the leaf-cutting ant Atta opaciceps,Borgmeier (Hymenoptera: Formicidae)

To understand the significance of the trail pheromone used in chemical communication of the leaf-cutting ants Atta opaciceps we investigated, under laboratory conditions, the trail-following behaviour of different castes. We observed a clear behavioural discrimination of conspecific venom gland extract of foraging ants from those of other species. Additionally, we determined the pheromone composition of A. opaciceps venom gland secretion using a two-dimensional gas chromatography coupled with mass spectrometry. Chemical analyses revealed the presence of three nitrogen-containing compounds, identified as 2,5-dimethylpyrazine, 3-ethyl-2,5-dimethylpyrazine and methyl 4-methylpyrrole-2-carboxylate (M4MPC). Four different bioassays performed with workers from different castes of A. opaciceps suggested that the trail pheromone elicits the trail-following behaviour in conspecifics of all castes, but the foragers respond more strongly to their own pheromone than to that of other castes (gardeners, generalists and soldiers). In addition, A. opaciceps foragers follow the trails made with the venom gland extracts of the unrelated Acromyrmex subterraneus subterraneus foragers as well as they follow the trails made with their own venom gland extract. M4MPC was identified to be the most abundant and the most behaviourally active component of the venom gland extract of A. opaciceps foragers.     

The Behavior of <Emphasis Type="Italic">Acromyrmex crassispinus</Emphasis> (Hymenoptera: Formicidae) on Trail Bifurcations and the Influence of Ant Flow on Error Rates of Nestbound Laden Workers

Ants are ordinarily faced with a succession of bifurcations along their foraging networks. Given that there is no directionality in pheromone trails, each bifurcation is potentially an opportunity for error in the trajectory of laden workers to the nest, which could entail considerable inefficiencies in the transportation of food to the colony. Leaf-cutting ants (Atta and Acromyrmex) commonly show intense traffic and complex foraging trail systems, which make them ideal organisms to study worker behavior in trail bifurcations. The behavior of leaf-cutting ants of the genus Acromyrmex in trail bifurcations is still largely unexplored. Thus, this study aimed to assess the behavior of Acromyrmex crassispinus workers on trail bifurcations and to investigate whether differences in ant flow on foraging trails influence the error rate of nestbound laden workers at trail bifurcation. There was a negative relationship between ant flow and error rate of nestbound laden workers. Most workers walked in the central part of the foraging trails but occupied a broader area of the foraging trail when the ant flow was high. The results of this study provide valuable insight into the organization of traffic flow in A. crassispinus and its impacts on the foraging strategy of the species.     

The trail of the african urticating antTetramorium aculeatum: Source,potency, and workers' behavior (Hymenoptera: Formicidae)

This paper studies the production of and the response to the trail in the African urticating ant,Tetramorium aculeatum under a variety of laboratory conditions. The trail was found to contain a complex mix of substances. Two of these components are secreted by the poison gland: The most volatile one is an attractant and increases the ants' linear speed; the other is the trail pheromone, which may act for days on a dry substrate. A third component is present on the last abdominal sternite. It acts as an attractant and a locostimulant and is synergistic of the trail pheromone. The activity of these substances increases with the age of the workers. While following a trail, foragers, even unrewarded, reinforce it with both the poison gland contents and the synergistic compound. The ants follow trails better in darkness than in light. A wetted trail rapidly loses its activity. The article suggests an explanation for the functioning ofT. aculeatum's natural trails, including the role of its different components.     

The Effect of Trail Pheromone and Path Confinement on Learning of Complex Routes in the Ant Lasius niger

Route learning is key to the survival of many central place foragers, such as bees and many ants. For ants which lay pheromone trails, the presence of a trail may act as an important source of information about whether an error has been made. The presence of trail pheromone has been demonstrated to support route learning, and the effect of pheromones on route choice have been reported to persist even after the pheromones have been removed. This could be explained in two ways: the pheromone may constrain the ants onto the correct route, thus preventing errors and aiding learning. Alternatively, the pheromones may act as a ‘reassurance’, signalling that the learner is on the right path and that learning the path is worthwhile. Here, we disentangle pheromone presence from route confinement in order to test these hypotheses, using the ant Lasius niger as a model. Unexpectedly, we did not find any evidence that pheromones support route learning. Indeed, there was no evidence that ants confined to the correct route learned at all. Thus, while we cannot support the ‘reassurance’ hypothesis, we can rule out the ‘confinement’ hypothesis. Other findings, such as a reduction in pheromone deposition in the presence of trail pheromones, are remarkably consistent with previous experiments. As previously reported, ants which make errors on their outward journey upregulate pheromone deposition on their return. Surprisingly, ants which would go on to make an error down-regulate pheromone deposition on their outward journey, hinting at a capacity for ants to gauge the quality of their own memories.     

Coordination of Raiding and Emigration in the Ponerine Army Ant Leptogenys distinguenda (Hymenoptera: Formicidae: Ponerinae): A Signal Analysis

Several glandular sources of trail pheromones have been discovered in army ants in general. Nevertheless, at present the understanding of the highly coordinated behavior of these ants is far from complete. The importance of trail pheromone communication for the coordination of raids and emigrations in the ponerine army ant Leptogenys distinguenda was examined, and its ecological function is discussed. The secretions of at least two glands organize the swarming activities of L. distinguenda. The pygidial gland is the source of an orientation pheromone holding the group of raiding workers together. The same pheromone guides emigrations to new nest sites. In addition, the poison sac contains two further components: one with a weak orientation effect and another which produces strong, but short-term attraction and excitement. The latter component is important in prey recruitment and characterizes raid trails. This highly volatile recruitment pheromone allows the extreme swarm dynamic characteristic of this species. Emigration trails lack the poison gland secretion. Due to their different chemical compositions, the ants are thus able to distinguish between raid and emigration trails. Nest emigration is not induced chemically, but mechanically, by the jerking movements of stimulating workers.     

Sensory Aspects of Trail-Following Behaviors in the Asian Longhorned Beetle, <Emphasis Type="Italic">Anoplophora glabripennis</Emphasis>

Anoplophora glabripennis has a complex suite of mate-finding behaviors, the functions of which are not entirely understood. These behaviors are elicited by a number of factors, including visual and chemical cues. Chemical cues include a male-produced volatile semiochemical acting as a long-range sex pheromone, a female-produced cuticular hydrocarbon blend serving as a sex-identification contact pheromone, and a recently identified female-produced trail sex pheromone that is followed by mate-seeking males. However, the sensory appendages and sensilla on these appendages used to detect the trail sex pheromone are unknown. We evaluated the ability of virgin male A. glabripennis to follow a sex pheromone trail after removal of the terminal four antennal segments and/or the maxillary and labial palps using a two-choice behavioral bioassay. We also tested the ability of males to follow the trail sex pheromone using volatile pheromone cues only, without physical contact with the pheromone. Results indicate that the palps are primarily responsible for sensing the pheromone, with males lacking palps unable to respond behaviorally to the trail sex pheromone. Under the conditions of this study, males could not follow the sex pheromone trail without direct contact, suggesting that olfaction may not be involved in detection of this pheromone. However, we did not determine to what degree the trail pheromone chemicals can volatilize under our experimental conditions. This work is important in elucidating the behaviors and sensory structures involved in mate-finding by this species on host trees, and these studies may help determine whether the trail sex pheromone has applications for monitoring and management.     

Toxicity, repellency, and transfer of chlorfenapyr against western subterranean termites (Isoptera: Rhinotermitidae)

Chlorfenapyr is a slow-acting insecticide against western subterranean termite, Reticulitermes hesperus Banks, when applied to sand. The LD50 at day 7 for workers is 29.98 ng per termite and considerably higher than that of chlorpyrifos (14.01), cypermethrin (3.21), and fipronil (0.16). Brief exposures to sand treated with chlorfenapyr resulted in dose-dependent mortality over a broad range of concentrations. Brief 1-h exposures to > or =75 ppm provided >88% kill of termites at day 7. Chlorfenapyr deposits did not repel termites, even at 300 ppm. Termites tunneled from 0.1 to 1.8 cm into sand treated with 10- to 300-ppm chlorfenapyr deposits, resulting in > or =70% mortality. Within 1 h after being exposed to 50 ppm chlorfenapyr, approximately 17% of the termites exhibited impaired responses to synthetic trail pheromone. By 4 h, nearly 60% of the workers were not able to follow a 10 fg/cm pheromone trail. There was a direct linear relationship of the uptake of [14C]chlorfenapyr as concentration and duration of exposure increased. The percentage of chlorfenapyr transferred to recipients varied from 13.3 to 38.4%. Donors exposed for 1 h transferred a greater percentage of chlorfenapyr than did donors exposed for 4 h. A 1-h exposure on 100-ppm deposits provided sufficient uptake to kill 100% of the donors and sufficient transfer to kill 96% of the recipients. There was not enough transfer for recipients to serve as secondary donors and kill other termites. Horizontal transfer is limited to contact with the original donor and by the decreased mobility of workers within 4-8 h after exposure to treated sand. The effectiveness of chlorfenapyr barrier treatments is primarily due to its nonrepellency and delayed toxicity.     

The sensory basis of trail following in some lepidopterous larvae: contact chemoreception

ABSTRACT. . Caterpillars of the genus Malacosoma follow trails of the chemical 5-beta-cholestane-3,24-dione, but nothing is known of how they perceive this compound, or more generally about the sensory basis of trail following in caterpillars. By selective ablations of chemosensory organs we show that, in Malacosoma , the trail chemical is perceived by the maxillary palpi. In another lepidopteran species, Yponomeuta cagnagellus , the palpi are needed to discriminate their own trails from a trail of Malacosoma. Malacosoma larvae also lose their specificity for conspecific trails when their palpi are ablated. Volatile cues evidently do not play a role in trail-following behaviour, since neither Malacosoma nor Yponomeuta can orient on a trail covered with fine nylon mesh. These data indicate that for Malacosoma , and probably also for Yponomeuta , contact chemoreception mediated by the maxillary palpi is the primary mode of pheromone perception. The evolution of receptor sensitivity to trail chemicals in caterpillars is discussed.     

Spitting out information: Trigona bees deposit saliva to signal resource locations

Stingless bees of the species Trigona spinipes (Fabricius 1793) use their saliva to lay scent trails communicating the location of profitable food sources. Extracts of the cephalic labial glands of the salivary system (not the mandibular glands, however) contain a large amount (approx. 74%) of octyl octanoate. This ester is also found on the scent-marked substrates at the feeding site. We demonstrate octyl octanoate to be a single compound pheromone which induces full trail following behaviour. The identification of the trail pheromone in this widely distributed bee makes it an ideal organism for studying the mechanism of trail following in a day flying insect.     

Better tired than lost: Turtle ant trail networks favor coherence over short edges

Creating a routing backbone is a fundamental problem in both biology and engineering. The routing backbone of the trail networks of arboreal turtle ants (Cephalotes goniodontus) connects many nests and food sources using trail pheromone deposited by ants as they walk. Unlike species that forage on the ground, the trail networks of arboreal ants are constrained by the vegetation. We examined what objectives the trail networks meet by comparing the observed ant trail networks with networks of random, hypothetical trail networks in the same surrounding vegetation and with trails optimized for four objectives: minimizing path length, minimizing average edge length, minimizing number of nodes, and minimizing opportunities to get lost. The ants’ trails minimized path length by minimizing the number of nodes traversed rather than choosing short edges. In addition, the ants’ trails reduced the opportunity for ants to get lost at each node, favoring nodes with 3D configurations most likely to be reinforced by pheromone. Thus, rather than finding the shortest edges, turtle ant trail networks take advantage of natural variation in the environment to favor coherence, keeping the ants together on the trails.     

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.     

Pheromone trail decay rates on different substrates in the Pharaoh's ant, Monomorium pharaonis

Abstract. Many ants use pheromone trails to organize collective foraging. This study investigated the rate at which a well‐established Pharaoh's ant, Monomorium pharaonis (L.), trail breaks down on two substrates (polycarbonate plastic, newspaper). Workers were allowed to feed on sucrose solution from a feeder 30 cm from the nest. Between the nest and the feeder, the trail had a Y‐shaped bifurcation. Initially, while recruiting to and exploiting the feeder, workers could only deposit pheromone on the branch leading to the feeder. Once the trail was established (by approximately 60 ants per min for 20 min), the ants were not allowed to reinforce the trail and were given a choice between the marked and unmarked branches. The numbers of ants choosing each branch were counted for 30 min. Initially, most went to the side on which pheromone had been deposited (80% and 70% on the plastic and paper substrates, respectively). However, this decayed to 50% within 25 min for plastic and 8 min for paper. From these data, the half‐life times of the pheromone are estimated as approximately 9 min and 3 min on plastic and paper, respectively. The results show that, for M. pharaonis, trail decay is rapid and is affected strongly by trail substrate.