Trail Marking and Processionary Behavior of the Larvae of the Weevil Phelypera distigma (Coleoptera: Curculionidae)

We present here the results of an investigation of the behavioral bases of the first documented instance of trail marking and processionary behavior in a beetle. The larvae of Phelypera distigma (Coleoptera: Curculionidae) forage communally, moving over the host plant in head-to-tail processions. Our study shows that the larvae secrete a pheromone from the ventral surface of the posterior abdomen that both elicits and guides the collective locomotion of the cohort. The pheromone is soluble in acetone and other nonpolar solvents and is relatively short-lived, eliciting trail following for less than 4 h after its deposition. When in processionary formations, larvae stimulate locomotion in others by rapidly bobbing their heads against sets of setae that occur on the lateral flanks of the posterior tips of the abdomens of precedent individuals. Larvae are also strongly attracted to tactile or chemotactile stimuli found at the tip of the abdomen of other larvae and their response to lures made of eviscerated abdomens show that such stimuli take precedence over the trail pheromone in eliciting and orienting locomotion. The cycloalexic formations adopted by resting larvae maximize the amount of body contact possible in a two-dimensional aggregate and allow tactile signals to rapidly radiate through the groups, alerting all members of a cohort to the onset of bouts of activity.     

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.     

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.     

Trail Marking and Abandonment of Depleted Feeding Sites by the Caterpillars of <Emphasis Type="Italic">Eutachyptera psidii</Emphasis> (Lepidoptera: Lasiocampidae)

The caterpillars of Eutachytptera psidii (Lepidoptera: Lasiocampidae) mark trails between their nest and distant feeding sites with a trail pheromone secreted from the ventral surfaces of their last abdominal segments. The threshold sensitivity to artificial trails prepared from an hexane extract of the pheromone was 0.75 × 10⁻³ caterpillar-equivalents per cm of trail. In tests of trail-specificity involving four other social species, the caterpillars responded only to trails prepared from a pheromone extract of the closely related genus Gloveria. Tests were conducted to determine the efficiency with which colonies abandon exhausted feeding sites in favor of new food finds. On their first forays after their food sources were experimentally moved from established feeding sites to new sites, 67.5 ± 3.6% of all the evening’s activity occurred on pathways leading to the previously established sites. During their next two forays, 86.3 ± 3.7% and 92.1 ± 2.0% of all activity occurred on pathways leading to the new sites. Efficient abandonment of exhausted feeding sites is attributed to the persistence of the trail pheromone, differential marking of new and old pathways and to the caterpillar’s ability to discern trail strength and to choose stronger over weaker trails at choice points.     

Trail following and stowaway behaviour of the myrmecophilous staphylinid beetle,Homoeusa acuminata,during foraging trips of its hostLasius fuliginosus (Hymenoptera: Formicidae)

Summary The behaviour of adultHomoeusa acuminata on trails of its hostLasius fuliginosus was investigated both in the field and in the laboratory. The beetles were active from May to September, accurately following the foraging trails of their hosts up to 20 metres from the nest. Most of the time, they were ignored by the ants, but if attacked they raised their abdomen as a possible appeasement or defensive behaviour. On trails the beetles most probably act as food robbers, feeding on prey collected by ants. The following method, called stowaway behaviour, was used by the beetles: when a beetle encountered an ant carrying a prey back to the nest it jumped on the prey, probably feeding on it while being transported.Laboratory experiments on circular artificial trails demonstrated thatH. acuminata follows a water extract of hindguts of the ants, the source of the trail pheromone. Both beetles and ants responded to an artificial trail of 0.03 hindgut equivalent per cm, but the mean distance followed by the beetles was about twelve times higher than that covered by the ants themselves. In contrast, experiments with solutions of the six fatty acids reported as the active components of the trail pheromone showed that the beetles did not respond at all, and that the ants only respond to the fatty acids at a very high concentration.     

Thigmotaxis maintains processions of late‐instar caterpillars of Ochrogaster lunifer

Processionary behaviour is a distinctive feature of the dispersal of caterpillars of a number of Lepidoptera, including Ochrogaster lunifer Herrich‐Schäffer (Lepidoptera: Notodontidae) from Australia. By cutting hairs on either the first or last three segments of the bodies of caterpillars and re‐introducing them into processions of normal caterpillars, the present study provides strong evidence that thigmotaxis is the mechanism that maintains single‐file, head‐to‐tail movement of groups of individuals. The inclusion of caterpillars whose posterior hairs have been cut results in the breakage of processions, whereas the inclusion of caterpillars whose anterior hairs have been cut rarely results in the breakage of processions. The breakage of processions occurs because caterpillars whose posterior hairs had been cut are difficult to follow and they will not stop when those behind became detached. The speed of movement of processions incorporating caterpillars whose hairs on the posterior segments of their bodies have been cut also has a significant effect on the likelihood of maintaining processionary movement; faster‐moving processions are more likely to break than are more slowly‐moving processions.     

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.     

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.     

Social Behavior of Larvae of the Neotropical Processionary Weevil Phelypera distigma (Boheman) (Coleoptera: Curculionidae: Hyperinae)

Socially gregarious behavior among free‐living leaf‐eating insect larvae occurs mostly among Lepidoptera, Symphyta, and a few Chyrsomelidae (Coleoptera). However, the Neotropical hyperine curculionid Phelypera distigma has also evolved this lifestyle, exhibiting a suite of social behaviors unique among beetles. The larvae are nomadic processionary foragers that punctuate foraging bouts with rosette‐shaped resting formations (cycloalexy). Larvae also vibrate or bob their heads rapidly when moving, especially when in contact with conspecifics, and this suggests acoustic or vibrational communication. In this study we used observational and experimental approaches to investigate the basis of processionary, cycloalexic,and head‐vibration behavior of this species. Larvae used both trail pheromones and thigmotactic signals to organize themselves into head‐to‐tail processionary columns. The trail pheromone, produced from the center of the abdomen, remains active for up to 4 h. Processions are not consistently led by particular individuals, but dynamically change over time and often temporarily break into two or more subprocessions. Subprocessions reunite through use of the trail pheromone. We found no evidence that head‐bobbing generates attraction through substrate‐borne or acoustic signals, but this behavior functions in direct contact to excite group activity. Time‐lapse videography used to analyze cycloalexic group formation showed that larvae transition from feeding in a line along the leaf margin to cycloalexic formations on the upper leaf surface via a coordinated back‐up movement that brings the posterior tip of their abdomens into contact. We identify three phases of cycloalexic formation: line‐up, back‐up, and an adjustment phase. Complete assembly can be achieved in as little as 5 s, but often the two phases establishing the basic rosette lasts 5–10 min, while the adjustment phase slowly tightens the group over a period of up to an hour. Collectively these studies present the first documented case of chemical trail marking in a beetle, and provide insight into a remarkable social‐behavioral repertoire convergent in key respects with the better‐studied social caterpillars and sawflies.     

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.

     

Utilization on extrafloral nectaries and fruit domatia of Canavalia lineata and C. cathartica (Leguminosae) by ants

Utilization of the extrafloral nectaries (EFNs) and fruits of Canavalia lineata and C. cathartica by ants was investigated at 30 sites in Japan. The fruits of C. lineata and C. cathartica were inhabited by five and eight ant species, respectively. Ant nesting periods and their utilization of EFNs differed between C. lineata and C. cathartica. Canavalia lineata flowers once a year, and periods of EFN-utilization and fruit-nesting by ants do not overlap. The fruit-nesting ants on C. lineata seem to invade the plant from the holes made by moth larvae or breaches made by decay. The ants nesting on the fruits of C. lineata may defend the plant against seed herbivores because they feed on moth larvae. Canavalia cathartica flowers several times over a year, and fruits are found throughout the year; therefore, periods of EFN-utilization and fruit-nesting by ants are overlapped. Canavalia cathartica offers year-round nesting sites and food for ants, and therefore may receive a higher defensive effect from ants than C. lineata. Handling editor: Graham Stone.     

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.     

Foraging behavior and growth of isolated larvae of a social caterpillar, Malacosoma americanum

In laboratory experiments, isolated eastern tent caterpillars, Malacosoma americanum (Fabricius) (Lepidoptera: Lasiocampidae), grew more slowly than their grouped siblings even though factors previously reported to give grouped caterpillars an advantage were eliminated from our experimental design. Analysis of time-lapse videorecordings of daily foraging bouts showed that, despite their slower growth, isolated individuals fed significantly more often than their grouped siblings. This finding is consistent with previous observations showing that the rate at which tent caterpillars assimilate food is largely independent of foraging frequency and suggests that increased metabolic costs associated with superfluous activity may cause isolated caterpillars to grow slower. More rapid growth of grouped caterpillars also appears attributable to the significantly longer periods of time solitary caterpillars spent inactive during episodes of molting. Our study shows that the distinctive temporal pattern of foraging characteristic of intact colonies of eastern tent caterpillars is a emergent property of the group.     

Territory defense by the ant Azteca trigona: maintenance of an arboreal ant mosaic

Mosaics of exclusive foraging territories, produced by intra-and interspecific competition, are commonly reported from arboreal ant communities throughout the tropics and appear to represent a recurring feature of community organization. This paper documents an ant mosaic within mangrove forests of Panama and examines the behavioral mechanisms by which one of the common species, Azteca trigona, maintains its territories. Most of the mangrove canopy is occupied by mutually exclusive territories of the ants A. trigona, A. velox, A. instabilis, and Crematogaster brevispinosa. When foraging workers of A. trigona detect workers of these territorial species, they organize an alarm recruitment response using pheromonal and tactile displays. Nestmates are attracted over short distances by an alarm pheromone originating in the pygidial gland and over longer distances by a trail pheromone produced by the Pavan's gland. Recruits are simultaneously alerted by a tactile display. No evidence was found for chemical marking of the territory. Major workers are proportionally more abundant at territory borders than on foraging trails in the interior of the colony. The mechanisms of territory defense in A. trigona are remarkably similar to those of ecologically analogous ants in the Old World tropics.     

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.     

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.     

The effect of seeds of exotic species transported via horse dung on vegetation along trail corridors

It has been suggested that exotic species will colonize within forests more frequently by the continual introduction of seeds through horse dung deposited along trails. Whether or not these exotic species have the ability to spread into and establish in the forest interior has been disputed. To address this, horse dung and soil samples were collected from trails during Autumn 1994 and Summer 1995 from three areas in southern Illinois, USA open to recreational horse travel. In addition, deer dung samples were collected from each of the study areas. Vegetation data were collected from each of the trail systems as well as from a trail along which horse travel was prohibited. The density of vascular plants in 0.25 m² quadrats placed at varying distances from the trail center to 5 m into the forest interior were recorded. Finally, dung samples were placed in situ along horse trails at one site to examine seedling germination in natural conditions. While 23 exotic species germinated from samples of horse dung placed out in a greenhouse, only one of these exotic species was also found in trail plots (Kummerowia striata). Similarly, while there were empirically more exotic species found along the trails allowing horse travel than there were on the trail lacking horse travel, the relative importance of those species was negligible along both trails. These results suggest that the emigration of exotic species via horse dung does not pose an immediate threat to the plant communities adjacent to trails in these forest systems. Nevertheless, the large number of exotic species in horse dung reflects the constant threat to any system from these species. Care must be taken, when allowing horseback use in areas, to anticipate invasion by exotic species from horse dung     

Silk drives aggregation and following in the neotropical caterpillar Mechanitis menapis (Nymphalidae: Ithomiini)

Gregarious larvae that use chemical communication to feed and move together are widespread among folivorous insects, although social behaviour has been studied almost exclusively in a few temperate zone genera. The Menapis (or variable) tigerwing butterfly Mechanitis menapis mantineus Hewitson (Lepidoptera, Nymphalidae, Danainae, Ithomiini) is a neotropical species whose larvae feed gregariously on Solanaceae host plants. In laboratory experiments conducted in the Ecuador cloud forest, M. menapis caterpillars are attracted to silk produced by conspecifics and show no evidence of pheromone production. Indeed, caterpillars consistently choose arenas with silk over bare arenas but do not show a preference for arenas marked with abdominal cuticular surface residues. Mechanitis menapis caterpillars on silk‐coated plants are both more mobile and more cohesive than those on control plants. Nonetheless, caterpillars move independently over unmarked surfaces and groups do not make rapid collective choices between two food sources. Collective behaviour in M. menapis thus appears to be based on aggregation on collectively produced silk to facilitate feeding, as well as using this silk to maintain cohesion. Silk production is common in caterpillars, although M. menapis appears to be unique among species studied so far in using silk to maintain group cohesion.     

The trail pheromone of the termite, Trinervitermes trinervoides

By using a T-junction choice assay workers of Trinervitermes trinervoides have been shown to lay and follow pheromone trails. The trials are not polarized and there is a quantitative relation between the number of termites laying the trail and the fraction of test termites which follw correctly. Workers reinforce trails whenever they follow them, and the degree of this reinforcement seems to be independent of trail strength. The trail pheromone is volatile and need not be perceived by contact chemoreception. Activity loss from filter paper is approximately exponential with a half-life of about 2 hr. Extracts of papers over which workers had laid trails were used to show a linear relationship, on a log-probit scale, between dose and response. Soldier termites also follow trails about as well as do workers, but workers lay trails that are about six to seven times as strong as soldier trails. No preference of either caste for their own trails could be demonstrated. Thin-layer chromatography indicated that the trail pheromone(s) is a fairly polar substance.     

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.