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Colonies of Pheidole ambigua ants excavate soil and drop it outside the nest entrance. The deposition of thousands of loads leads to the formation of regular ring-shaped piles. How is this pattern generated? This study investigated soil pile formation on level and sloping surfaces, both empirically and using an agent-based model. We found that ants drop soil preferentially in the direction in which the slope is least steeply uphill from the nest entrance, both when adding to an existing pile and when starting a new pile. Ants respond to cues from local slope to choose downhill directions. Ants walking on a slope increase the frequency and magnitude of changes in direction, and more of these changes of direction take them downhill than uphill. Also, ants carrying soil on a slope wait longer before dropping their soil compared to ants on a level plane. These mechanisms combine to focus soil dropping in the downhill direction, without the necessity of a direct relationship between slope and probability of dropping soil. These empirically determined rules were used to simulate soil disposal. The slight preference for turning downhill measured empirically was shown in the model to be sufficient to generate biologically realistic patterns of soil dumping when combined with memory of the direction of previous trips. From simple rules governing individual behaviour an overall pattern emerges, which is appropriate to the environment and allows a rapid response to changes. \ud

The struggle among social classes or castes is well known in humans. Here, we show that caste inequality similarly affects societies of ants, bees and wasps, where castes are morphologically distinct and workers have greatly reduced reproductive potential compared with queens. In social insects, an individual normally has no control over its own fate, whether queen or worker, as this is socially determined during rearing. Here, for the first time, we quantify a strategy for overcoming social control. In the stingless bee Schwarziana quadripunctata, some individuals reared in worker cells avoid a worker fate by developing into fully functional dwarf queens. \ud \ud

In human society selfish use of common resources\ud can lead to disaster, a situation known as the ‘tragedy\ud of the commons’ (TOC). Although a TOC is\ud usually prevented by coercion, theory predicts that\ud close kinship ties can also favour reduced exploitation.\ud We test this prediction using data on a TOC\ud occurring in Melipona bee societies.

We investigated worker policing by egg eating in the ponerine ant Pachycondyla inversa, a species with\ud morphologically distinct queens and workers. Colonies were split into one half with the queen and one\ud half without. Workers in queenless colony fragments started laying unfertilized male eggs after three weeks.\ud Worker-laid eggs and queen-laid eggs were introduced into five other queenright colonies with a single\ud queen and three colonies with multiple queens, and their fate was observed for 30 min. Significantly more\ud worker-laid eggs (range of 35–62%, mean of 46%) than queen-laid eggs (range of 5–31%, mean of 15%)\ud were eaten by workers in single-queen colonies, and the same trend was seen in multiple-queen colonies.\ud This seems to be the first well-documented study of ants with a distinct caste polymorphism to show that\ud workers kill worker-laid eggs in preference to queen-laid eggs. Chemical analyses showed that the surfaces\ud of queen-laid and worker-laid eggs have different chemical profiles as a result of different relative proportions\ud of several hydrocarbons. Such differences might provide the information necessary for differential\ud treatment of eggs. One particular alkane, 3,11-dimeC27, was significantly more abundant on the surfaces\ud of queen-laid eggs. This substance is also the most abundant compound on the cuticles of egg layers.

Magnetic resonance imaging (MRI) enables in vivo imaging of organisms. The recent development of the magnetic resonance microscope (MRM) has enabled organisms within the size range of many insects to be imaged. Here, we introduce the principles of MRI and MRM and review their use in entomology. We show that MRM has been successfully applied in studies of parasitology, development, metabolism, biomagnetism and morphology, and the advantages and disadvantages relative to other imaging techniques are discussed. In addition, we illustrate the images that can be obtained using MRM. We conclude that although MRM has significant potential, further improvements to the technique are still desirable if it is to become a mainstream imaging technology in entomology. Abbreviation: CSI chemical shift imaging. The dependence of the resonance frequency of a nucleus on the chemical binding of the atom or molecule in which it is contained. (N)MRI (nuclear) magnetic resonance imaging MRM magnetic resonance microscopy Voxel A contraction for volume element, which is the basic unit of MR reconstruction; represented as a pixel in the display of the MR image.

Animal societies are stages for both conflict and cooperation. Reproduction is often monopolized by one or a few individuals who behave aggressively to prevent subordinates from reproducing (for example, naked mole-rats, wasps and ants). Here we report an unusual mechanism by which the dominant individual maintains reproductive control. In the queenless ant Dinoponera quadriceps, only the alpha female reproduces. If the alpha is challenged by another female she chemically marks the pretender who is then punished by low-ranking females. This cooperation between alpha and low-rankers allows the alpha to inflict punishment indirectly, thereby maintaining her reproductive primacy without having to fight

Mutual policing is thought to be important in conflict suppression at all levels of biological organization. In hymenopteran societies (bees, ants, and wasps), multiple mating by queens favors mutual policing of male production among workers (worker policing). However, worker policing of male production is proving to be more widespread than predicted by relatedness patterns, occurring in societies headed by single-mated queens in which, paradoxically, workers are more related to the workers' sons that they kill than the queen's sons that they spare. Here we develop an inclusive-fitness model to show that a second reproductive conflict, the conflict over sex allocation, can explain the evolution of worker policing contrary to relatedness predictions. Among ants, and probably other social Hymenoptera, workers kill males to favor their more related sisters. Importantly, males are killed at the larval stage, presumably because workers cannot determine the sex of queen-laid eggs. Sex-allocation biasing favors worker policing because policing removes some males (the workers' sons) at low cost at the egg stage rather than at higher cost at the larval stage. Our model reveals an important interaction between two reproductive conflicts in which the presence of one conflict (sex allocation) favors the suppression of the other (male production by workers).\ud \ud

The collection and handling of colony resources such as food, water, and nest-construction material is often divided into subtasks in which the material is passed from one worker to another. This is known as task partitioning. If task; are partitioned with direct transfer of material between foragers and receivers, queueing delays can occur as individuals search or wait for a transfer partner. Changes in environmental conditions and relative number of foragers and receivers affect these delays as well as colony ergonomic efficiency. These delays are used in recruitment in both honeybees and Polybia wasps. This study investigates the distribution of queueing delays and the information content and quality of those delays using a stochastic-simulation model. Information quality increases with colony size. When the relative proportions of foragers and receivers are suboptimal, the group in excess has better information. Individuals can increase information quality of delays by two mechanisms: averaging over consecutive trips and averaging over multiple transfers within a trip where direct transfer occurs. We suggest that multiple transfer occurs in the honeybee in order to improve information quality. \ud

The collection and handling of colony resources such as food, water, and nest construction material is often divided into subtasks in which the material is passed from one worker to another. This is known as task partitioning. When material is transferred directly from one individual to another, queueing delays frequently occur because individuals must sometimes wait for a transfer partner. A stochastic simulation model was written to study the effect of colony size on these delays. Queueing delay decreases roughly exponentially with colony size because stochastic fluctuations in the arrival of individuals are lower in larger colonies. These results support empirical studies of Polybia occidentalis and other theoretical studies of honeybees. The effect of the relative number of individuals in the two subtask groups was also studied. There is a unique optimal ratio of the number of workers associated with each of the subtasks that simultaneously minimizes mean queueing delay and maximizes colony nectar-processing rate. Deviations from this optimal ratio, for example, as a result of forager mortality or changes in nectar productivity that affect foraging trip duration, increase mean queueing delays greatly, especially in smaller colonies.\ud