The trail following behaviour of Yponomeuta cagnagellus

In this study the trail following behaviour of the caterpillar Yponomeuta cagnagellus (Hübner) (Lepidoptera, Yponomeutidae) is investigated. It is demonstrated that these caterpillars follow trails made by conspecifics. Ablation experiments show that at least part of this behaviour is directed by the tactile senses, but additional chemical cues cannot be excluded. In choice experiments using trails from different species, Y. cagnagellus strongly preferred conspecific trails over those from Malacosoma neustria, but did not prefer conspecific over other Yponomeuta trails. This lack of species-specificity within Yponomeuta is discussed and it is concluded that trail following is probably of little help in the elucidation of the evolutionary history of the genus.

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La façon dont Yponomeuta cagnagellus suit les pistes

Résumé Cette étude traite de la façon dont les chenilles de Y. cagnagellus Hübner (Lépido. Yponomeutidae) suivent des pistes. Elles suivent les pistes de leurs congénères. Par ablation, on peut montrer qu'une partie au moins de ce comportement est due à des perceptions tactiles, mais que l'intervention d'informations chimiques complémentaires ne peut être exclue. En présence de choix entre traces de différentes espèces, elles préfèrent nettement les traces des congénères à celles de Malacosoma neustria, mais ne choisissent pas entre les différentes espèces d'yponomeutes. L'absence de spécificité parmi le genre Yponomeuta montre que la sélection des pistes est probablement de peu d'utilité pour l'étude de l'histoire évolutive du genre.

     

Social Cues and Following Behavior in the Forest Tent Caterpillar

Colonies of the social caterpillar Malacosoma disstria Hubner (Lepidoptera: Lasiocampidae) travel in groups following silk trails marked with pher-omone. This study examined first, the cues involved in following behavior and second, the responses to these cues at different larval stadia. Both second and fourth instar larvae discriminated between fresh and older trails, and travelled faster in the presence of trails. In addition to trail following, young caterpillars exhibited leader following, which might be particularly important in exploring unmarked territory. Indeed, second instar caterpillars were more likely to travel together when trails were absent. Fourth instar larvae exhibited greater independent locomotion in the absence of trails than did younger larvae. These findings help explain patterns of social behavior observed in forest tent caterpillar colonies in the field.     

Host specificity of trail marking to foliage by eastern tent caterpillars, Malacosoma americanum

The recruitment trail marking behavior of eastern tent caterpillars (Malacosoma americanum Fabr.) was modified by rearing them on plants which they do not usually attack in nature. Caterpillars reared on one of two nonhosts (Prunus avium (L.) L. or Quercus coccinea Muenchh.) marked pheromone trails to foliage of their rearing plant, whereas caterpillars reared on a natural host plant (Prunus serotina Ehrh.) did not mark trails to nonhost foliage. Caterpillars preferred host to nonhost foliage, regardless of their rearing history. The degree of trail marking was correlated with suitability of foliage for larval growth. The results indicate that trail marking behavior is a response to relative rather than absolute food quality, but that preference behavior is more rigidly programmed to favor the optimal food.

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Zusammenfassung Das Rekrutierungs-Spurmarkierungsverhalten von Malacosoma americanum Fabr. wurde durch die Zucht der Raupen auf Nichtwirtspflanzen modifiziert. Die Bevorzugung der Wirtspflanze jedoch wurde nicht verändert. Das Spurmarkierungsverhalten wurde im Laboratorium quantifiziert, indem die Zahl der markierten Abschnitte eines Kartonstreifens gezählt wurde, auf dem die Raupe von einem Ende zum andern kroch. Das Präferenzverhalten wurde geprüft, indem Blattscheiben von Wirts-und Nichtwirtspflanzen an einem Ende des Kartonstreifens befestigt wurden. Die Raupen markierten Spuren zu Nichtwirtspflanzen (Prunus avium oder Quercus coccinea) nur, wenn sie darauf gezüchtet worden waren. Andererseits war die Spurmarkierung auf Wirtspflanzen (Prunus serotina) hin intensiv unabhängigig von der Art der Zucht. Ebenfalls unabhängig von der Aufzucht war die Bevorzugung von P. serotina vor Nichtwirtspflanzen in Wahlversuchen. Präferenz und Spurmarkierung waren korreliert mit der Eignung der Blätter für Raupenwachstum. Die Resultate zeigen, dass das Spurmarkierungsverhalten mehr eine Reaktion auf relative als auf absolute Futterqualität ist, dass jedoch Präferenz strenger auf optimales Futter programmiert ist.

     

Trail-Based Communication in Social Caterpillars of Eriogaster lanestris (Lepidoptera: Lasiocampidae)

Caterpillars of Eriogaster lanestris (Lepidoptera: Lasiocampidae) mark trails as they move between feeding sites and their communal tent. They prefer new trails over aged ones. Hungry caterpillars prefer trails marked by fed caterpillars returning to the tent. Thus successful foragers direct tentmates to profitable food finds, in a manner similar to that of Malacosoma americanum. E. lanestris readily follows trails prepared from 5-cholestane-3-one, a component of the trail marker of M. americanum, when applied at a rate of 10^–10 g/cm. In choice tests, they preferred more highly concentrated over weaker trails. New trails are always established in groups, and the velocity increases from early to late caterpillars traversing a new trail.     

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.     

Communication of leaf suitability by gregarious eastern tent caterpillars (Malacosoma americanum)

ABSTRACT. 1. Previous work has shown that leaf age affects recruitment trail marking by eastern tent caterpillars ( Malacosoma americanum Fabr.). Young leaves of host plants elicit trail marking to a greater degree than mature leaves.
2. Experiments were conducted to establish the relationship between the differential behavioural responses of larvae to young and mature leaves and the suitability of foliage for larval growth and survival. Foliage of black cherry ( Prunus serotina Ehrh.), a typical rosaceous host plant, was used for this comparison.
3. Larvae preferred young leaves to mature leaves in choice tests, and marked more to young leaves than to mature leaves in no-choice tests.
4. Mature leaves supported adequate growth through two larval instars of rearing, but thereafter were unsuitable for growth. Larvae fed mature leaves had lower pupal weight, poorer survival, and grew less efficiently than larvae fed young leaves.
5. The results support the hypothesis that the trail communication system of eastern tent caterpillars is an adaptation to efficiently locate leaves which are favourable for larval growth and survival.     

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.     

Do trails affect relative abundance estimates of rainforest frogs and lizards?

The selection of a sampling protocol is critical to study amphibian and reptile communities and in many instances researchers have combined the use of visual encounter surveys (VES) conducted on trails and off trails. The effect of human‐made trails on relative abundance estimates of amphibians and reptiles has been assessed in a few temperate locations, but data are lacking for tropical sites. Our study was designed to address this issue by comparing abundance estimates of frogs and lizards on and off trails in a lowland rainforest in south‐eastern Perú. We used nocturnal VES to sample frogs and lizards along transects established on trails and off trails in two different forest types. We found that the observed relative abundance estimates of frogs and lizards were affected by the location of transects (on trail vs. off trail) and the type of forest (floodplain forest vs. terra firme forest). We also found an interaction between the two main effects, indicating that the effect of transect location with respect to trails varies as a function of habitat. Observed frog abundances were higher on trails than off trails, indicating that studies that include VES on trails will bias relative abundance estimates in contrast to studies that include only VES off trails. We suggest that transects should be established only off trails, especially for monitoring studies because trail use by humans can have a strong influence on observed animal abundance.     

Trail Marking by the Larva of the Madrone Butterfly Eucheira socialis and the Role of the Trail Pheromone in Communal Foraging Behavior

The larva of the Madrone butterfly Eucheira socialis (Lepidoptera: Pieridae) secretes a trail pheromone from the ventral surface of the posterior tip of its abdomen. Caterpillars mark trails by bringing the secretory site into brief contact with the substrate during a locomotive cycle. Foragers mark most heavily when they move onto new branches and little, if at all, when they move over established trails or when they return to the communal shelter after feeding. The caterpillars make careful comparisons of alternative pathways at choice points and select newer and stronger trails over older and weaker trails. Differential marking of new and established trails during nightly forays, coupled with sensory discrimination of trails by strength and age, leads colonies to abandon old trails in favor of new trails. When applied at a rate as low as 2.5 × 10 ^–10 g/mm, caterpillars followed synthetic trails prepared from 5-cholestane-3-one, a trail pheromone previously reported from the tent caterpillars (Malacosoma spp.). Although both Eucheira and Malacosoma mark with the tip of the abdomen and have near-identical sensitivites to 5-cholestane-3-one, our study shows that Eucheira employs a relatively unsophisticated system of trail-based communication and does not recruit to food. The trail-based communication system of Eucheira appears to represent an early stage in the evolution of cooperative foraging that is derived from, and motivationally linked to, conflict behavior.     

Maintenance of foraging trails by the giant tropical antParaponera clavata (Insecta: Formicidae: Ponerinae)

Summary Establishment and maintenance of foraging trails to an artificial nectar source by ten colonies ofParaponera davata (Fabr.) in Panama is reported. The first forager to locate the artificial nectar source was responsible for recruiting additional foragers and for marking trails to orient these foragers. More than half of the trail marking was performed by the first two ants to mark the path back to the colony, although up to 11 ants per colony per hour marked trails. The number of trail marks and the number of marking ants decreased through time, presumably as foragers learned the location of the artificial nectar source. Four categories of recruits were noted: markers, foragers, patrollers, and visitors.     

Chemical trail systems,orientation, and territorial interactions in the antLasius neoniger

Foraging and territoriality in the ant Lasius neonigerinvolves a series of trails which channel foragers away from adjacent colonies. Experimental studies suggest that the trails are composed of colony-specific, persistent orientation components of hindgut material that accumulate on trails during foraging. A less durable component of the hindgut trail pheromone regulates recruitment. Foraging directionality and the use of a trail could be modified by experimentally arranging confrontations with conspecifics. The orientation of foragers is mediated by visual as well as chemical cues. Components of the foraging and territorial system of L. neonigerappear to include (1) a network of subnests which change in position seasonally within each polydomous nest; (2) a series of trails emanating from each subnest that adjusts search toward resource patches and away from aggressive, neighboring conspecifics; and (3) trail communication involving an ephemeral component of the hindgut trail pheromone that regulates the organization of cooperative prey retrieval and a more persistent component that serves as an orientation guide.     

Differential Use of Trails by Forest Mammals and the Implications for Camera-Trap Studies: A Case Study from Belize

Relative abundance indices are often used to compare species abundance between sites. The indices assume that species have similar detection probabilities, or that differences between detection probabilities are known and can be corrected for. Indices often consist of encounter frequencies of footprints, burrows, markings or photo captures along trails or transect lines, but the assumption of equal detection probabilities is rarely validated. This study analyzes detection probabilities of a range of Neotropical mammals on trails in dense secondary forests, using camera-trap and track data. Photo captures of the two large cats, jaguars ( Panthera onca ) and pumas ( Puma concolor ), were correlated solely with trail variables, while photo captures of their potential prey species had no correlation or negative correlation with trail variables. The Neotropical mammals varied greatly in their tendency to follow or cross trails based on footprints surveys. This indicates that camera locations on trails will have varying detection probability for these Neotropical mammals. Even the two similar-sized jaguars and pumas, occupying relatively similar niches, differed subtly in their use of trails. Pumas followed trails more completely while jaguars were more likely to deviate from trails. The ecological significance of these findings is that jaguars seem to be more willing to use the forest matrix away from trails than do pumas. We conclude that trail-based indices, such as photographic captures or tracks along trails, are not appropriate for comparison between Neotropical species, and not even between relatively similar species like jaguars and pumas.     

Host-ant trail following by myrmecophilous larvae of Liphyrinae (Lepidoptera,Lycaenidae)

In this study we report a case of ant-trail following by lycaenid caterpillars. Euliphyra mirifica and E. leucyana caterpillars are involved in a commensal association with the weaver ant Oecophylla longinoda. The host nests are made with leaves which over the course of time dry out or are broken open by storms, forcing the ants to migrate and build a new nest elsewhere. Euliphyra caterpillars are stimulated by recruitment behaviour which triggers the migration of their host. They then follow the host trails leading to the new nesting site. Laboratory experiments showed that these caterpillars are able to follow host trails under varied conditions: (1) fresh trails actually used by workers, (2) fresh trails in the absence of workers, (3) heterocolonial, 2-month-old trails, and (4) fresh trails washed with water (to simulate the effect of tropical rains). They can also bridge trail gaps of more than 1 cm. Under natural conditions, the trails are frequently situated along thin twigs. The forward progress of the ants in such a situation is not impeded by the presence of large Euliphyra larvae. Workers just climb over the caterpillars, even on larger trails where there is enough room to pass alongside them. This suggests that an allomone is secreted on the dorsal part of the caterpillars. When crawling along heterocolonial trails, the caterpillars are not attacked, even if about 21% of the workers from the new colony spread their mandibles when encountering them. They are then adopted and are admitted to the nest of the new host colony of O. longinoda.     

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.     

Simple models for trail-following behaviour; Trunk trails versus individual foragers

Social insects such as ants use trial-marking and trail-following to organize the behaviour and movement patterns of a large population. Since behaviour has to meet needs of the population in a changing environment, the type of trail networks formed must be adaptable. Both solitary foraging as well as mass migration along a system of trunk trails are behaviours essential for survival of the colony, and the population must be able to switch from one behaviour to the other, depending on conditions. Using a mathematical model for trail following we show that subtle changes in individual behaviour can give rise to dramatic differences in the behaviour of the population, including the ability to switch from solitary movement to organized group traffic. The model incorporates biological parameters associated with the organism, the trail-marker, and the population. Ordinary differential equations are formulated for the density of the trails and for the number of individuals following trails or exploring randomly. It is assumed that the followers reinforce trails by pheromone marking, and that individuals respond to the strength of the trails by becoming more efficient followers. The model is analyzed by qualitative phase-plane methods.     

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.     

An agent-based model to investigate the roles of attractive and repellent pheromones in ant decision making during foraging

Pharaoh's ants organise their foraging system using three types of trail pheromone. All previous foraging models based on specific ant foraging systems have assumed that only a single attractive pheromone is used. Here we present an agent-based model based on trail choice at a trail bifurcation within the foraging trail network of a Pharaoh's ant colony which includes both attractive (positive) and repellent (negative) trail pheromones. Experiments have previously shown that Pharaoh's ants use both types of pheromone. We investigate how the repellent pheromone affects trail choice and foraging success in our simulated foraging system. We find that both the repellent and attractive pheromones have a role in trail choice, and that the repellent pheromone prevents random fluctuations which could otherwise lead to a positive feedback loop causing the colony to concentrate its foraging on the unrewarding trail. An emergent feature of the model is a high level of variability in the level of repellent pheromone on the unrewarding branch. This is caused by the repellent pheromone exerting negative feedback on its own deposition. We also investigate the dynamic situation where the location of the food is changed after foraging trails are established. We find that the repellent pheromone has a key role in enabling the colony to refocus the foraging effort to the new location. Our results show that having a repellent pheromone is adaptive, as it increases the robustness and flexibility of the colony's overall foraging response.     

Effects of gaster extract trail concentration on the trail following behaviour of the Argentine ant, Iridomyrmex humilis (Mayr)

In the Argentine ant, optimum trail following to gaster extracts was displayed to 0.1 and 1.0 equivalents/50 cm of circular trail. Trail following to airborne components was demonstrated when ants exhibited normal trail following behaviour while walking 3 or 6 mm below a 0.1 ant equivalent trail. However, at 8 or 12 mm separation, following ceased, indicating that the height of the active space was ca. 6–8 mm. The average horizontal distance from the centre of the trail at which ants exhibited following behaviour increased with concentration to 3–4 mm beyond the applied trail boundaries, indicating both an ability to follow airborne chemicals, and possibly a non-tolerance of excessively high concentration. Activity of 0.1 ant equivalent trails on filter paper declined to about half the original level by four hours; after eight hours, responses were significantly different from, but almost as low as, solvent controls.     

Adaptative properties of the chemical trail system of the African weaver antOecophylla longinoda Latreille (Hymenoptera,Formicidae, Formicinae)

Summary Trail communication of the weaver antOecophylla longinoda is highly adapted to the African rain forests, insofar natural selection in the tropics might favour chemical trails which are durable in nature. A dry chemical trail can last over nine weeks, and over ten months when reinforced with fecal marking. The trail pheromones are resistant to rain, whether they are fresh or three months old, and whether they are reinforced or not with anal deposits.     

The Role of Tactile and Chemical Stimuli in the Formation and Maintenance of the Processions of the Social Caterpillar Hylesia lineata (Lepidoptera: Saturniidae)

Colonies of the social caterpillar Hylesia lineata (Lepidoptera: Satumiidae) form long, single-file, head-to-tail processions as they move between their shelters and distant feeding sites. Although investigations of other processionary species have implicated a silk trail in the processionary process, silk plays little or no role in initiating or maintaining processions in H. lineata. Studies we report here implicate both tactile stimuli and a trail pheromone in the establishment and maintenance of processions. Processionaries elicit locomotion in the individual preceding them in line by brushing their heads against prominent sulci that project from the tips of their abdomens. Caterpillars mark their pathways with a pheromone deposited by brushing the ventral surfaces of their last abdominal segments against the substrate. The persistent pheromone is soluble in hexanes and appears to be secreted from glandular setae found on the proximal regions of the anal prolegs and the venter. In Y-choice tests, caterpillars selected newer trails over older trails and stronger trails over weaker trails. They did not distinguish between trials deposited by newly fed caterpillars and those deposited by starved caterpillars. Despite the unidirectional nature of processions, there is no indication that caterpillars can determine from the trail alone the direction in which the procession advanced. The significance of these findings to the foraging ecology of the caterpillars is discussed.