Communal Foraging Behavior and Recruitment Communication in Gloveria sp.

The caterpillars of Gloveria sp. mark trails with a pheromone they deposit by dragging the ventral surface of the tip of the abdomen along branch pathways as they move between their communal nest and distance feeding sites. The threshold sensitivity of the caterpillar for an extract prepared from the secretory site was approximately 0.5 × 10^–3 caterpillar equivalents/cm of trail. Bioassays show that Gloveria follows neither authentic trails of Malacosoma americanum nor artificial trails prepared from 5-cholestane-3-one, a chemical previously reported to elicit trail following from other social caterpillars. Although our observations show that fed caterpillars mark heavily as they return to their nest, we found no evidence that individual caterpillars are able to recruit hungry nestmates to new food finds. In this species, recruitment to food occurs only after many caterpillars have reinforced a trail to a newly discovered food source. In contrast, hungry caterpillars of the confamilial species M. americanum, tested under identical conditions, responded strongly to the postprandial trails of individual caterpillars and rapidly abandoned depleted sites in favor of new food finds. We postulate that the difference in the efficiency with which these two species recruit nestmates to food evolved in response to differences in the spatial distribution of their food supplies.     

Behavioral evidence for multicomponent trail pheromone in the termite,Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae)

Evidence is presented for a multicomponent trail pheromone in the eastern subterranean termiteReticulitermes flavipes (Kollar). Choice tests were used to compare strength and persistency of trails made by termites in glass tubes. Tubes connected to termite nests for 24 hr were marked by termites with an extremely long-lasting chemical that persisted for at least one year, and a highly volatile substance that decayed in 15 min. Individual termites varied the trail they deposited under different circumstances. When removed from the nest and placed in a clean tube, they deposited a highly volatile substance. They left stronger and more persistent trails when exploring clean tubes attached to the nest. When they discovered a new food source, they left trails that were more attractive and far more persistent than trails made by exploring or displaced termites. A food trail made by 15 workers was still effective after 24 hr, whereas a trail made by 50 displaced workers, walking one after another through a tube, lasted only 5 min.     

Specificity of trail markers of forest and eastern tent caterpillars

Exploratory trails deposited on paper strips by the forest tent caterpillar (FTC),Malacosoma disstria Hubner, and the eastern tent caterpillar (ETC),M. americanum (Fabricius), as well as extracts of these trails, readily elicited interspecific trail-following behavior. In 2-choice tests involving simple Y mazes constructed from these paper strips, the caterpillars of both species preferred by approximately 31 the trails of the FTC. Studies involving whole colonies of the ETC maintained under nearnatural conditions in the laboratory, however, indicated that the trails deposited by successful foragers of the ETC as they returned to their tent from feeding sites were more attractive than the exploratory trails of either the ETC or FTC. The pronounced interspecific response of these congeners to each other's trails suggests that they utilize either qualitatively similar or identical trail-marking chemicals. Both species preferred their own trails to those ofArchips cerasivoranus (Fitch) (Tortricidae), providing the first evidence that more distantly related lepidopterous larvae utilize distinct trails.     

A chemical trail factor from the silk of the eastern tent caterpillarMalacosoma americanum (Lepidoptera: Lasiocampidae)

The leaf-feeding larvae of the eastern tent caterpillarMalacosoma americanum (Fabricius) follow silk trails laid down on branches leading from their communal tent to distant foraging sites. The response of colonies reared in the laboratory under seminatural conditions to silk trails washed in methylene chloride and to chemical trails prepared from a solvent extract of their tent or trail silk, showed that one or more soluble components of their trail is essential to the elicitation of the following response. The demonstrated ability of the caterpillars to distinguish between old and newly reinforced silk trails most likely occurs in response to a temporal change in the detectable chemical properties of their trail.     

Chemical trail marking and following by caterpillars ofMalacosoma neustria

Chemical trail marking and following by gregarious caterpillars,Malacosoma neustria L. (Lepidoptera: Lasiocampidae), was studied in the laboratory. As in other species ofMalacosoma, larvae deposit a trail pheromone from a sternal secretory site when searching the host for food. Larvae in the vanguard of foraging columns establish chemical trails as they explore new territory. Marking behavior diminishes as successive unfed foragers utilize the trail. These exploratory trails are subsequently overmarked by fed larvae returning to the tent. Other foragers follow the trails of fed larvae in preference to trails of unfed larvae. Thus, like the eastern tent caterpillar,M. americanum, successful foragers ofM. neustria recruit colony-mates to feeding sites. The chemical activity of both recruitment and exploratory trails degrades slowly, suggesting that the trail pheromone ofM. neustria is a nonvolatile substance. Caterpillars ofM. neustria readily follow the nonvolatile trail pheromone which has been identified fromM. americanum, 5-cholestane-3,24-dione.     

Hexyl Decanoate, the First Trail Pheromone Compound Identified in a Stingless Bee, Trigona recursa

Foragers of many species of stingless bees guide their nestmates to food sources by means of scent trails deposited on solid substrates between the food and the nest. The corresponding trail pheromones are generally believed to be produced in the mandibular glands, although definitive experimental proof has never been provided. We tested the trail following behavior of recruits of Trigona recursa in field experiments with artificial scent trails branching off from natural scent trails of this stingless bee. First-time recruits (newcomers) did not follow these trails when they were laid with pure solvent or mandibular gland extract. However, they did follow trails made with labial gland extract. Chemical analyses of labial gland secretions revealed that hexyl decanoate was the dominant component (72.4 ± 1.9% of all volatiles). Newcomers were significantly attracted to artificial trails made with synthetic hexyl decanoate, demonstrating its key function in eliciting scent-following behavior. According to our experiments with T. recursa, the trail pheromone is produced in the labial glands and not in the mandibular glands. Hexyl decanoate is the first component of a trail pheromone identified and proved to be behaviorally active in stingless bees.     

Trail and arena marking by caterpillars ofArchips cerasivoranus (lepidoptera: Tortricidae)

The activity ofArchips cerasivoranus caterpillars is largely limited to their colonial silk web and trails. Silk pulled directly from the spinnerets of caterpillars and wound onto paper strips to form artificial trails elicited locomotion from the larvae. Trails made from extracts of silk and silk glands also elicited locomotion. These and other observations reported here indicate that the caterpillars are responsive to a water-soluble pheromone that is a component of the silk strand. Marker pheromones appear not to be secreted from other regions of the body, as has been reported for some other trail-following caterpillars.     

Behavioral bioassays of termite trail pheromones

The monocyclic 14-membered ring diterpene, cembrene-A, previously identified as a nasutitermitine trail pheromone, was tested for its effectiveness as a trail pheromone inNasutitermes costalis. Artificial trails prepared from serial dilutions of racemic cembrene-A over a concentration range of 10^–1–10^–6 mg/ml were ineffective in recruiting termites. Serial dilutions of racemic cembrene-A ranging in concentration from 10^–1 to 10^–5 mg/ml produced an orientation effect. Chiral cembrene-A produced recruitment in soldiers at 10^–1 and 10^–3 mg/ml and was less ineffective in recruiting workers. Soldiers always showed a lower and more variable recruitment response to chiral cembrene-A than to sternal gland extracts. The behavioral response to both chiral and racemic cembrene-A was different in quantity and quality from that observed for sternal gland extract. Based on the results of these behavioral tests, cembrene-A appears to be a generalized nasute orientation pheromone which may show recruitment properties at unnaturally high concentrations.     

Trail-following behavior ofReticulitermes hesperus Banks (Isoptera: Rhinotermitidae)

The behavior ofReticulitermes hesperus Banks pseudergates (workers) was assessed on artificial trails containing different concentrations of sternal gland extract. On nongiadient trails, more pseudergates were recruited to trails of greater pheromone concentration, they traveled a greater distance without pausing, and their rate of locomotion increased over that observed on trails of lesser concentration (positive orthokinesis). Of the individuals pausing before completing trails of high concentration, fewer left the trails or reversed direction (negative klinokinesis) than on trails of lower concentration. Termites walking down concentration gradients failed to complete these trails to the low-concentration termini. At a point representing an average decrease of slightly more than 10-fold in the original concentration of pheromone, individuals reversed their direction of travel and returned to the high-concentration terminus. Termites walking up pheromone gradients proceeded to the high-concentration termini without reversing direction.R. hesperus pseudergates are thus able to orient along a gradient of trail pheromone by longitudinal klinotaxis.     

Chemoorientation of eastern tent caterpillars to trail pheromone, 5β-Cholestane-3,24-dione

Chemoorientation behavior of the larval eastern tent caterpillar,Malacosoma americanum, was studied using the synthetic trail pheromone 5-cholestane-3,24-dione. Divergent arms of Y mazes were treated with various concentration ratios of the pheromone. At application rates of 10^–10-10^–9 g/mm of trail, larvae showed a significant preference for stronger trails when concentration ratios differed by as little as 4:1. At application rates of 10^–8 and greater there was no significant difference in trail choice even when trails differed in strength by a full order of magnitude. Other studies showed that the caterpillars abandon the pattern of choosing stronger over weaker trails when they repeatedly fail to find food at the end of a stronger trail. Experiments in which larvae were required to choose trails separated by a gap demonstrated orientation by chemoklinotaxis. Caterpillars that had one of the maxillary palps ablated looped in the direction of their intact chemo-receptor when placed on filter paper treated uniformly with pheromone, indicating that they may also orient by tropotaxis. The relevance of these findings to the tent caterpillar communication system is discussed.     

Isolation of the trail recruitment pheromone ofSolenopsis invicta

TheSolenopsis invicta trail pheromone is synthesized by the Dufour's gland and is released through the sting apparatus. The recruitment subcategory of theS. invicta trail pheromone was shown to be composed of a mixture of the orientation pheromone, (Z,E)--farnesene and an unidentified homosesquiterpene consisting of three rings and one double bond (C-1). C-1 is present in worker Dufour's glands at only 75 pg per worker equivalent. This is the first report that demonstrates that different exocrine products from the same gland control different subcategories of behavior related to mass recruitment.     

Chemical communication in the primitive antAneuretus simoni: The role of the sternal and pygidial glands

Trail and alarm communication inAneuretus simoni are mediated by the secretions of the sternal and pygidial glands, respectively. The sternal gland is composed of a glandular epithelium and an associated reservoir located in the 7th sternum. This gland produces a relatively long-lived mass recruitment pheromone. The pygidial gland opens between the 6th and 7th tergites and produces a secretion that releases aggressive alarm. The evolution of glandular physiology in the Aneuretinae and Dolichoderinae are discussed in view of these findings.     

Chemical marker from silk ofYponomeuta cagnagellus

Trail following in lepidopterous larvae is often attributed to chemical markers, but only a few clear-cut examples are found in the literature. In this paper evidence is presented for a chemical basis of the trail following behaviour ofYponomeuta cagnagellus. (Lepidoptera: Yponomeutidae) The marker is shown to be very persistent under laboratory conditions and is water soluble. Several possible secretory sites were investigated, and it is concluded that the marker is probably secreted together with the silk from the labial gland. Problems associated with the demonstration of trail markers in caterpillars are discussed.     

Role of trail pheromone in foraging and processionary behavior of pine processionary caterpillars Thaumetopoea pityocampa

Although caterpillars of Thaumetopoea pityocampa may mark their pathways with silk, this study shows that the material is essential to neither the elicitation nor maintenance of trail-following or processionary behavior. Trail following is dependent upon a pheromone the caterpillars deposit by brushing the ventral surfaces of the tips of their abdomens against the substrate. Earlier instars are strongly bound to their trail system; in the laboratory, caterpillars followed circular trails continuously for as long as 12 hr before breaking away from them. The trail pheromone is long-lived and soluble in nonpolar solvents, but its volatilization or degradation allows the caterpillars to distinguish new from aged trails. In contrast to trail following, processionary behavior, the head-to-tail, single-file movement of the caterpillars is dependent on neither silk nor the trail pheromone. Stimuli associated with setae found on the tip of the abdomen of the precedent caterpillar serve to hold processions together, and such stimuli take priority over those associated with either the trail pheromone or silk. Although the caterpillars discern trail strength and choose stronger over weaker trails, the trail marking system of the processionary caterpillar appears less sophisticated than those of other, previously studied species of social caterpillars, and colonies are relatively inefficient in abandoning exhausted feeding sites in favor of new food finds. In laboratory studies, females were more likely to lead processions than males, and leaders, regardless of gender, expended more energy in locomotion than followers.     

Trail following and recruitment: Response of eastern tent caterpillarMalacosoma americanum to 5β-cholestane-3,24-dione and 5β-cholestan-3-one

Studies were conducted to determine the relative effectiveness of 5-cholestane-3,24-dione (diketone) and 5-cholestan-3-one (monoketone) in eliciting trail following from eastern tent caterpillars,Malacosoma americanum. In Y maze tests, trails prepared from the monoketone were followed preferentially over diketone trails, even when the diketone trail was several orders of magnitude stronger. Under field conditions, colonies readily abandoned well-developed trail systems in favor of artificial trails that were established with the monoketone. Other tests in which the caterpillars selected trails prepared from the monoketone (but not the diketone) more often than their own recruitment trails indicate that the monoketone constitutes the chemical basis of recruitment communication in this insect. The study also shows that tent caterpillars are highly sensitive to small differences in the amount of monoketone in a trail and can distinguish between new and aged trails prepared from the compound.     

Scent trailing by virgin females ofPseudococcm calceolariae

Virgin females of the citrophilous mealybugPseudococcus calceolariae (Mask.) deposit scent marks as trails on the substrate on which they rest or move. These substances elicit attraction and sexual behavior by conspecific males. The same responses were obtained when males were bioassayed on extracts from filter paper disks on which females had rested. The significance of scent trailing in mealybugs is discussed.Homoptera, Coccoidea, Pseudococcidae.This work was supported by the Italian National Research Council (CNR).     

Conspecific scent trailing by garter snakes (Thamnophis sirtalis) during autumn Further evidence for use of pheromones in den location

Adult garter snakes (Thamnophis sirtalis), collected in October near a traditional, communal hibernaculum in central Wisconsin, were acclimated to autumnal conditions and subjected to laboratory tests to determine whether they could follow scent trails of a conspecific. Graded responses were obtained, but 75% of the sample showed at least some inclination to follow scent trails. The results suggest that pheromone cues may be used by male and female garter snakes to locate traditional dens during autumnal migrations. Evidence from this and other studies suggests that pheromones are probably used in conjunction with other homing mechanisms and that the role of pheromones in den location may be more important in younger snakes and in populations inhabiting northern latitudes.     

Trail-following in termites: Evidence for a multicomponent system

Several African termite species from different subfamilies and different habitats are sensitive to trail-active extracts or to naturally laid trails from other species. Using single-extract bioassays, it is shown that the response threshold for trail-following is nearly identical for all tested species (except forHodotermes mossambicus). However, when termite workers have a choice between trails from their own species and from other species, conspecific trail-following is exclusively observed. This phenomenon can be counteracted by dilution (110) of the conspecific trail-pheromone extract. Tests of the trail activity of various synthetic alcohols show that among these, the highest sensitivity of termite workers is to (Z)-3-dodecen-1-ol. Based on our experimental data, we postulate that, in addition to a generally active trail-pheromone constituent (an unsaturated primary C₁₂ alcohol) or a pool of chemically closely related alcohols, other species-specific components are present in termite trails.     

Role of (3<Emphasis Type="Italic">Z</Emphasis>,6<Emphasis Type="Italic">Z</Emphasis>,8<Emphasis Type="Italic">E</Emphasis>)-Dodecatrien-1-ol in Trail Following,Feeding, and Mating Behavior of <Emphasis Type="Italic">Reticulitermes hesperus</Emphasis>

Trail pheromones mediate communication among western subterranean termites, Reticulitermes hesperus Banks. Repetitive passages of ≥28 termites were required to establish a pheromone trail and trails needed to be reinforced because they lasted <48 hr. The minimal threshold concentration for inducing responses from termite workers and secondary reproductives was between 0.01 and 0.1 fg/cm of (3Z,6Z,8E)-dodecatrien-1-ol (henceforth, dodecatrienol). Workers showed optimal trail-following behavior to dodecatrienol at a concentration of 10 fg/cm. Trails with concentrations >10 pg/cm were repellent to workers. Workers did not detect pheromone gradients, responding equally to increasing or decreasing gradients of dodecatrienol, and termite workers were not able to differentiate between different concentrations of dodecatrienol. Termites preferred dodecatrienol trails to 2-phenoxyethanol trails. Antennae played a key role in trail pheromone perception. Dodecatrienol acted as an arrestant for worker termites (10 fg/cm²) and male alates (5 ng/cm²), whereas sternal gland extracts from females attracted male alates. Workers and alates, upon contact with filter paper disks treated with higher doses (10 fg/cm² and 5 ng/cm², respectively) of dodecatrienol, were highly excited (increased antennation and palpation) and repeatedly returned to the treated disks. Dodecatrienol did not act as a phagostimulant when offered on a paper towel disk. Reticulitermes hesperus is highly responsive to dodecatrienol, and it may play an important role in orientation of workers and alates.     

Chemical cues used by prairie rattlesnakes (Crotalus viridis) to follow trails of rodent prey

Each of 10 prairie rattlesnakes (Crotalus viridis) was exposed to three types of trails after striking rodent prey (Mus musculus). One trail was made with mouse urine, another was made with tap water, and the third consisted of materials from mouse integument. The snakes exhibited trailing behavior only when integumentary trails were available. It was concluded that prairie rattlesnakes do not utilize urinary cues; instead they attend to materials associated with rodent skin and fur.