Destruction of Spiderwebs and Rescue of Ensnared Nestmates by a Granivorous Desert Ant (Veromessor pergandei).

Prey species rarely seek out and dismantle traps constructed by their predators. In the current study, we report an instance of targeted trap destruction by an invertebrate and a novel context for rescue behavior. We found that foragers of the granivorous desert ant (Veromessor pergandei) identify and cooperatively dismantle spiderwebs (Araneae: Theridiidae, Steatoda spp., and Asagena sp.) During group foraging, workers ensnared in webs are recovered by sisters, which transport them to the nest and groom away their silk bindings. The presence of an ensnared nestmate and chemical alarm signal significantly increased the probability of web removal and nestmate retrieval. A subset of larger-bodied foragers participated in web removal, and 6.3% became tangled or were captured by spiders. Most animals that perform rescue behavior live in small groups, but V. pergandei colonies include tens of thousands of short-lived workers. To maintain their size, large colonies must collect enough seeds to produce 650 new ants each day. We hypothesize that the removal of spiderwebs allows for an unimpeded income of seeds on a single foraging path during a brief daily temperature window. Despite the cost to individuals, webs are recognized and removed only when workers are captured in them.

Larval regulation of worker reproduction in the polydomous ant Novomessor cockerelli.

Although workers in many ant species are capable of producing their own offspring, they generally rear the queen's offspring instead. There are various mechanisms that regulate worker reproduction including inhibitory effects of ant brood. Colonies of the ant Novomessor cockerelli are monogynous and polydomous resulting in a large portion of nest workers being physically isolated from the queen for extended periods of time. Some workers experimentally isolated from the queen in laboratory nests lay viable eggs, which develop into males. We investigate the mechanism that regulates worker fertility in subnests separated from the queen by giving queenless worker groups queen-produced larvae, queen-produced eggs, or no brood. Our findings show that larvae delay the time to worker egg-laying, but eggs have no effect. Larval inhibition is a likely mechanism that contributes to the regulation of worker reproduction in N. cockerellli because larvae are easily transported to subnests that do not contain a queen.

Chemical communication during foraging in the harvesting ants Messor pergandei and Messor andrei.

We combined behavioral analyses in the laboratory and field to investigate chemical communication in the formation of foraging columns in two Nearctic seed harvesting ants, Messor pergandei and Messor andrei. We demonstrate that both species use poison gland secretions to lay recruitment trails. In M. pergandei, the recruitment effect of the poison gland is enhanced by adding pygidial gland secretions. The poison glands of both species contain 1-phenyl ethanol. Minute quantities (3 µl of a 0.1 ppm solution) of 1-phenyl ethanol drawn out along a 40 cm long trail released trail following behavior in M. pergandei, while M. andrei required higher concentrations (0.5-1 ppm). Messor pergandei workers showed weak trail following to 5 ppm trails of the pyrazines 2,5-dimethylpyrazine and 2,3,5-trimethylpyrazine, whereas M. andrei workers showed no behavioral response. Minute quantities of pyrazines were detected in M. pergandei but not in M. andrei poison glands using single ion monitoring gas chromatography-mass spectrometry.

A context-dependent alarm signal in the ant Temnothorax rugatulus.

Because collective cognition emerges from local signaling among group members, deciphering communication systems is crucial to understanding the underlying mechanisms. Alarm signals are widespread in the social insects and can elicit a variety of behavioral responses to danger, but the functional plasticity of these signals has not been well studied. Here we report an alarm pheromone in the ant Temnothorax rugatulus that elicits two different behaviors depending on context. When an ant was tethered inside an unfamiliar nest site and unable to move freely, she released a pheromone from her mandibular gland that signaled other ants to reject this nest as a potential new home, presumably to avoid potential danger. When the same pheromone was presented near the ants' home nest, they were instead attracted to it, presumably to respond to a threat to the colony. We used coupled gas chromatography/mass spectrometry to identify candidate compounds from the mandibular gland and tested each one in a nest choice bioassay. We found that 2,5-dimethylpyrazine was sufficient to induce rejection of a marked new nest and also to attract ants when released at the home nest. This is the first detailed investigation of chemical communication in the leptothoracine ants. We discuss the possibility that this pheromone's deterrent function can improve an emigrating colony's nest site selection performance.

An empirically based simulation of group foraging in the harvesting ant, Messor pergandei.

We present an empirically based group model of foraging interactions in Messor pergandei, the Sonoran desert harvesting ant. M. pergandei colonies send out daily foraging columns consisting of tens of thousands of individual ants. Each day, the directions of the columns may change depending on the resource availability and the neighbor interactions. If neighboring columns meet, ants fight, and subsequent foraging is suppressed. M. pergandei colonies face a general problem which is present in many systems: dynamic spatial partitioning in a constantly changing environment, while simultaneously minimizing negative competitive interactions with multiple neighbors. Our simulation model of a population of column foragers is spatially explicit and includes neighbor interactions. We study how different behavioral strategies influence resource exploitation and space use for different nest distributions and densities. Column foraging in M. pergandei is adapted to the spatial and temporal properties of their natural habitat. Resource and space use is maximized both at the colony and the population level by a model with a behavioral strategy including learning and fast forgetting rates.

New exocrine glands in ants: the hypostomal gland and basitarsal gland in the genus Melissotarsus (Hymenoptera: Formicidae).

Fisher and Robertson (Insect Soc 46: 78-83, 1999) discovered the production of silk-like secretions emerging from slit-shaped openings along the anterior margin of the ventral hypostoma of Melissotarsus ant workers. The current histological study describes a hitherto unknown hypostomal gland from which this silk-like substance originates. In addition, this study describes a new basitarsal gland in the three pairs of legs of Melissotarsus workers.

Worker division of labor and endocrine physiology are associated in the harvester ant, Pogonomyrmex californicus.

In Pogonomyrmex californicus harvester ants, an age-associated division of labor occurs in the worker caste, in which young workers perform in-nest tasks and older workers forage for food. Here, we tested whether this behavioral division is age based or age flexible, and whether it coincides with differential expression of systemic hormones with known roles in behavioral regulation. Whole-body content of juvenile hormone (JH) and ecdysteroids was determined in workers from (1) age-typical colonies, in which a typical age structure is maintained and workers transition across behaviors naturally, and (2) single-cohort colonies, which are entirely composed of same-aged workers, facilitating the establishment of age-independent division of labor. Foragers from both colony types had higher JH and lower ecdysteroid content than workers performing in-nest tasks, suggesting that age is not the sole determinant of worker behavior. This association between hormone content and behavior of P. californicus workers is similar to that previously observed in founding queens of this species. Because these hormones are key regulators of development and reproductive behavior, our data are consistent with the reproductive ground plan hypothesis (RGPH), which posits that the reproductive regulatory mechanisms of solitary ancestors were co-opted to regulate worker behavior.

Reclaiming the crown: queen to worker conflict over reproduction in Aphaenogaster cockerelli.

In many social taxa, reproductively dominant individuals sometimes use aggression to secure and maintain reproductive status. In the social insects, queen aggression towards subordinate individuals or workers has been documented and is predicted to occur only in species with a small colony size and a low level of queen-worker dimorphism. We report queen aggression towards reproductive workers in the ant species Aphaenogaster cockerelli, a species with a relatively large colony size and a high level of reproductive dimorphism. Through analysis of cuticular hydrocarbon profiles, we show that queens are aggressive only to reproductively active workers. Non-reproductive workers treated with a hydrocarbon typical for reproductives are attacked by workers but not by queens, which suggests different ways of recognition. We provide possible explanations of why queen aggression is observed in this species.

Behavior and exocrine glands in the myrmecophilous beetle Dinarda dentata (Gravenhorst, 1806) (Coleoptera: Staphylinidae: Aleocharinae).

The nests of advanced eusocial ant species can be considered ecological islands with a diversity of ecological niches inhabited by not only the ants and their brood, but also a multitude of other organisms adapted to particular niches. In the current paper, we describe the myrmecophilous behavior and the exocrine glands that enable the staphylinid beetle Dinarda dentata to live closely with its host ants Formica sanguinea. We confirm previous anecdotal descriptions of the beetle's ability to snatch regurgitated food from ants that arrive with a full crop in the peripheral nest chambers, and describe how the beetle is able to appease its host ants and dull initial aggression in the ants.

The non-additive effects of body size on nest architecture in a polymorphic ant.

Like traditional organisms, eusocial insect societies express traits that are the target of natural selection. Variation at the colony level emerges from the combined attributes of thousands of workers and may yield characteristics not predicted from individual phenotypes. By manipulating the ratios of worker types, the basis of complex, colony-level traits can be reduced to the additive and non-additive interactions of their component parts. In this study, we investigated the independent and synergistic effects of body size on nest architecture in a seasonally polymorphic harvester ant, Veromessor pergandei Using network analysis, we compared wax casts of nests, and found that mixed-size groups built longer nests, excavated more sand and produced greater architectural complexity than single-sized worker groups. The nests built by polymorphic groups were not only larger in absolute terms, but larger than expected based on the combined contributions of both size classes in isolation. In effect, the interactions of different worker types yielded a colony-level trait that was not predicted from the sum of its parts. In nature, V. pergandei colonies with fewer fathers produce smaller workers each summer, and produce more workers annually. Because body size is linked to multiple colony-level traits, our findings demonstrate how selection acting on one characteristic, like mating frequency, could also shape unrelated characteristics, like nest architecture.This article is part of the theme issue 'Interdisciplinary approaches for uncovering the impacts of architecture on collective behaviour'.

Behavior and exocrine glands in the myrmecophilous beetle Lomechusoides strumosus (Fabricius, 1775) (formerly called Lomechusa strumosa) (Coleoptera: Staphylinidae: Aleocharinae).

To become integrated into an ant society, myrmecophilous parasites must overcome both the defenses and the communication system of their hosts. Some aleocharine staphylinid beetles employ chemical and tactile strategies to invade colonies, where they later consume ant brood and participate in parasitic trophallaxis with host ants. By producing compounds that both appease their hosts and stimulate adoption, the beetles are able to live in and deposit their own eggs in the well defended ant nest. In the current paper, previous findings on the myrmecophilous behavior and morphological features of the staphylinid beetle Lomechusoides (formerly Lomechusa) strumosus are reviewed and re-evaluated. Hitherto unpublished results concerning the beetles' ability to participate in the social food flow of their host ants are reported. Furthermore, we present an analysis and documentation of the behavioral interactions between beetles and host ants during the adoption process, and we report new histological and scanning electron microscopic analyses of the exocrine glands and morphological adaptations that underlie the myrmecophilous behavior of L. strumosus. The main features of L. strumosus are compared with those of the staphilinid myrmecophile Lomechusa (formerly Atemeles) pubicollis. The paper concludes with a description of the life trajectory of L. strumosus and presents a brief history and discussion of the hypotheses concerning the evolution of myrmecophily in L. strumosus and other highly adapted myrmecophilous parasites.

Amphotis marginata (Coleoptera: Nitidulidae) a highwayman of the ant Lasius fuliginosus.

The space occupied by evolutionarily advanced ant societies can be subdivided into functional sites, such as broodchambers; peripheral nest chambers; kitchen middens; and foraging routes. Many predators and social parasites are specially adapted to make their living inside specific niches created by ants. In particular, the foraging paths of certain ant species are frequented by predatory and kleptoparasitic arthropods, including one striking example, the nitidulid beetle, Amphotis marginata. Adults of this species obtain the majority of their nutrition by acting as a kind of "highwayman" on the foraging trails of the ant Lasius fuliginosus, where they solicit regurgitation from food laden ant-workers by mimicking the ant's food-begging signals. Employing food labeled with the radio isotope 32P, we assessed the quantities of food the beetles siphoned-off of food-laden ants, and we investigated the site preferences, behavioral mechanisms and possible morphological adaptations underlying the food kleptoparasitism of A. marginata.

Queen Specific Exocrine Glands in Legionary Ants and Their Possible Function in Sexual Selection.

The colonies of army ants and some other legionary ant species have single, permanently wingless queens with massive post petioles and large gasters. Such highly modified queens are called dichthadiigynes. This paper presents the unusually rich exocrine gland endowment of dichthadiigynes, which is not found in queens of other ant species. It has been suggested these kinds of glands produce secretions that attract and maintain worker retinues around queens, especially during migration. However, large worker retinues also occur in non-legionary species whose queens do not have such an exuberance of exocrine glands. We argue and present evidence in support of our previously proposed hypothesis that the enormous outfit of exocrine glands found in dichthadiigynes is due to sexual selection mediated by workers as the main selecting agents.

Biosynthesis of the insect pheromone (S)-4-methyl-3-heptanone.

Using stable isotope-labelled probes and mass spectrometry, the insect pheromone (S)-4-methyl-3-heptanone is shown to be biosynthesised from three propionate units following a polyketide/fatty acid-type metabolic route.

Food robbing in ants, a form of interference competition.

Food robbing is a special form of interference competition in ants. It has been frequently observed in Myrmecocystus mimicus, which waylay returning foragers of several Pogonomyrmex species at their nests and take insect prey, particularly termites, away from them. Myrmecocystus more successfully robs prey from P. desertorum and P. maricopa than from P. barbatus. Usually only those Pogonomyrmex nests are affected by Myrmecocystus prey robbing that are not farther away than 10 m from the Myrmecocystus nest. Some M. mimicus workers show a high specialization in prey robbing. Other cases of food robbing in ants are described.

Interference strategy of Iridomyrmex pruinosum (Hymenoptera: Formicidae) during foraging.

The dolichoderine ant Iridomyrmex pruinosum and the formicines Myrmecocystus mimicus and M. depilis overlap widely in their niches. Although a worker of I. pruinosum is about four times smaller than a forager of Myrmecocystus, Iridomyrmex usually succeeds in displacing Myrmecocystus from the bait and often successfully blocks the nest entrance of its competitors, thus preventing Myrmecocystus foragers from leaving their nest. Iridomyrmex achieves this by quickly channeling large numbers of workers to food sources and nest entrances of Myrmecocystus, employing an effective chemical mass recruitment system, and chemicall repelling its competitors with secretions from the pygidial (anal) gland.

[Chemical strategy during foraging in Solenopsis fugax Latr. and Monomorium pharaonis L.] / Chemische Strategie beim Nahrungserwerb der Diebsameise (Solenopsis fugax Latr.) und der Pharaoameise (Monomorium pharaonis L.).

1. Solenopsis fugax, a subterranean ant, lays odorous trails in the tunnels leading to the brood chambers of neighbouring ant species. The recruitment trail pheromone originates from the Dufour's gland. 2. In addition, when preying on the foreign brood, Solenopsis fugax discharges a highly effective and long lasting repellent substance from the poison gland. This prevents the brood keeping ants from defending their own larvae against the predators. 3. A very similar chemical foraging strategy is used by Monomorium pharaonis. A repellent substance enabling Monomorium to compete successfully with other ant species for food sources is discharged. The recruitment pheromone originates from the Dufour's gland; a repellent substance enabling Monomorium to compete successfully with other ant species is discharged from the poison gland.

Leaf density and a trade-off between load-size selection and recruitment behavior in the ant Atta cephalotes.

This study considers the interplay between individual load-size selection and recruitment behavior in the leaf-cutting ant Atta cephalotes. Foraging workers anchor themselves on the leaf edge by their hind legs and pivot around them while cutting arcs from leaves. Since workers not only cut leaves but also lay chemical trails to recruit nestmates, we investigated whether there is conflict of motivation affecting the workers' decision either to quickly inform nestmates about a newly discovered food source, or instead to cut full-load leaf fragments, which could delay recruitment. Workers were presented with leaves of privet of three different grades of toughness (measured as leaf density=mass/area) as sources of different quality, and load-size selection and recruitment behavior by harvesting-satiated and harvesting-deprived workers were measured. The following results were obtained. (1) Leaf density affected individual load-size selection: both harvesting-satiated and harvesting-deprived workers were found to cut smaller leaf fragments from the denser leaves. (2) Harvesting-deprived workers cut smaller fragments than harvesting-satiated workers, and therefore saved cutting time. The fragments cut were smaller only during the initial phases of the recruitment process, when information about the discovery needed to be transferred. (3) Harvesting-deprived workers showed higher recruitment rates than harvesting-satiated workers. A considerable number of ants were observed to return to the nest unladen. During the initial phases, the ratio of laden/unladen workers was lower than that for harvesting-satiated workers, and increased with the development of the tograging process. (4) Scout workers confronted with familiar leaves ran back to the nest laying chemical trails without even contacting the leaves. They relied on olfactory cues to start recruiting nestmates, and leaf density played no role in their decisions. (5) When confronted with unfamiliar leaves, on the other hand, they assessed leaf quality by probing bites at the leaf edge, although no actual cuts occurred. In this situation, the resulting recruitment rates depended on physical leaf traits, being higher for the tenderer leaves. (6) Workers foraging on unfamiliar leaves cut smaller fragments than workers cutting familiar leaves, and most of them displayed trail-laying behavior when returning to the nest. The results support the hypothesis of a trade-off between time spent collecting and that invested to recruit nestmates. During the initial phases of exploitation of a newly discovered food source, workers reduced their individual carrying performance in order to return earlier to the colony for further recruitment.

Ants andPolyommatus icarus immatures (Lycaenidae) -sex-related developmental benefits and costs of ant attendance.

Third and fourth instar larvae and pupae of the facultatively myrmecophilous Palaearctic blue butterflyPolyommatus icarus showed no alteration in developmental time when reared in the presence of two species ofLasius ants. Sex differences were observed in larval growth and pupal weight, with males growing larger and faster. Sex-related differences also occurred in the costs and benefits of ant-attendance. Male pupal masses tended to be larger in individuals associated with ants, and their pupal weight loss was not enhanced by ant attendance. This positive developmental effect of myrmecophily is tentatively attributed to a stimulating influence of ants on caterpillar feeding behavior. In contrast, females associated with ants tended to lose more weight during the pupal stage. Hence there is evidence for developmental benefits, rather than costs, of myrmecophily in maleP. icarus immatures, whereas ant attendance appears to be more costly for females during the pupal stage. These findings are discussed in relation to data on other myrmecophilous lycaenid species. It is suggested that maintaining low-level myrmecophily and its related organs is a comparatively inexpensive evolutionary stable strategy among this butterfly group.