Path formation in a robot swarm

We present two swarm intelligence control mechanisms used for distributed robot path formation. In the first, the robots form linear chains. We study three variants of robot chains, which vary in the degree of motion allowed to the chain structure. The second mechanism is called vectorfield. In this case, the robots form a pattern that globally indicates the direction towards a goal or home location. We test each controller on a task that consists in forming a path between two objects which an individual robot cannot perceive simultaneously. Our simulation experiments show promising results. All the controllers are able to form paths in complex obstacle environments and exhibit very good scalability, robustness, and fault tolerance characteristics. Additionally, we observe that chains perform better for small robot group sizes, while vectorfield performs better for large groups.     

Invasive interactions: can Argentine ants indirectly increase the reproductive output of a weed?

The direct and indirect interactions of invasive ants with plants, insect herbivores, and Hemiptera are complex. While ant and Hemiptera interactions with native plants have been well studied, the effects of invasive ant–scale insect mutualisms on the reproductive output of invasive weeds have not. The study system consisted of Argentine ants (Linepithema humile), boneseed (Chrysanthemoides monilifera monilifera), and sap-sucking scale insects (Hemiptera: Saissetia oleae and Parasaissetia nigra), all of which are invasive in New Zealand. We examined the direct and indirect effects of Argentine ants on scale insects and other invertebrates (especially herbivores) and on plant reproductive output. Argentine ants spent one-third of their time specifically associated with scale insects in tending behaviours. The invertebrate community was significantly different between uninfested and infested plants, with fewer predators and herbivores on ant-infested plants. Herbivore damage was significantly reduced on plants with Argentine ants, but sooty mould colonisation was greater where ants were present. Herbivore damage increased when ants were excluded from plants. Boneseed plants infested with Argentine ants produced significantly more fruits than plants without ants. The increase in reproductive output in the presence of ants may be due to increased pollination as the result of pollinators being forced to relocate frequently to avoid attack by ants, resulting in an increase in pollen transfer and higher fruit/seed set. The consequences of Argentine ant invasion can be varied; not only does their invasion have consequences for maintaining biodiversity, ant invasion may also affect weed and pest management strategies.     

The evolution of plant cultivation by ants

Outside humans, true agriculture was previously thought to be restricted to social insects farming fungus. However, obligate farming of plants by ants was recently discovered in Fiji, prompting a re-examination of plant cultivation by ants. Here, we generate a database of plant cultivation by ants, identify three main types, and show that these interactions evolved primarily for shelter rather than food. We find that plant cultivation evolved at least 65 times independently for crops (~200 plant species), and 15 times in farmer lineages (~37 ant taxa) in the Neotropics and Asia/Australasia. Because of their high evolutionary replication, and variation in partner dependence, these systems are powerful models to unveil the steps in the evolution and ecology of insect agriculture.     

Evolving coordinated group behaviours through maximisation of mean mutual information

A well known problem in the design of the control system for a swarm of robots concerns the definition of suitable individual rules that result in the desired coordinated behaviour. A possible solution to this problem is given by the automatic synthesis of the individual controllers through evolutionary or learning processes. These processes offer the possibility to freely search the space of the possible solutions for a given task, under the guidance of a user-defined utility function. Nonetheless, there exist no general principles to follow in the definition of such a utility function in order to reward coordinated group behaviours. As a consequence, task dependent functions must be devised each time a new coordination problem is under study. In this paper, we propose the use of measures developed in Information Theory as task-independent, implicit utility functions. We present two experiments in which three robots are trained to produce generic coordinated behaviours. Each robot is provided with rich sensory and motor apparatus, which can be exploited to explore the environment and to communicate with other robots. We show how coordinated behaviours can be synthesised through a simple evolutionary process. The only criteria used to evaluate the performance of the robotic group is the estimate of mutual information between the motor states of the robots.     

Evolution of influence: signaling in a lycaenid-ant interaction

Some phytophagous insects gain defense from natural enemies by associating with otherwise potentially harmful top predators. Many lycaenid butterfly caterpillars are involved in such interactions with ants: larvae provide carbohydrate rewards from the dorsal nectary organ (DNO) to associated ants in return for protection from natural enemies. The stability of these interactions involves signals that identify the lycaenid caterpillar as a mutualist. However, larvae of some lycaenid species, such as Lycaena xanthoides, are found in close association with ants but do not possess the reward producing DNO. Evaluating the relationship in a phylogenetic framework, we show that the association between L. xanthoides and ants likely evolved from a non-ant-associated ancestor. Behavioral trials also show that L. xanthoides larvae are capable of influencing ant behavior to increase ant tending when faced with a simulated predator attack, without providing DNO-derived rewards to ant associates. These results demonstrate that the DNO is not necessary to maintain associations between lycaenid larvae and ants. Third-party interactions may affect the evolution of mutualisms and consideration of underlying evolutionary history is necessary to understand contemporary species associations.     

Corpse Management in Social Insects

Undertaking behavior is an essential adaptation to social life that is critical for colony hygiene in enclosed nests. Social insects dispose of dead individuals in various fashions to prevent further contact between corpses and living members in a colony. Focusing on three groups of eusocial insects (bees, ants, and termites) in two phylogenetically distant orders (Hymenoptera and Isoptera), we review mechanisms of death recognition, convergent and divergent behavioral responses toward dead individuals, and undertaking task allocation from the perspective of division of labor. Distinctly different solutions (e.g., corpse removal, burial and cannibalism) have evolved, independently, in the holometabolous hymenopterans and hemimetabolous isopterans toward the same problem of corpse management. In addition, issues which can lead to a better understanding of the roles that undertaking behavior has played in the evolution of eusociality are discussed.     

Death of an order: a comprehensive molecular phylogenetic study confirms that termites are eusocial cockroaches

Termites are instantly recognizable mound-builders and house-eaters: their complex social lifestyles have made them incredibly successful throughout the tropics. Although known as 'white ants', they are not ants and their relationships with other insects remain unclear. Our molecular phylogenetic analyses, the most comprehensive yet attempted, show that termites are social cockroaches, no longer meriting being classified as a separate order (Isoptera) from the cockroaches (Blattodea). Instead, we propose that they should be treated as a family (Termitidae) of cockroaches. It is surprising to find that a group of wood-feeding cockroaches has evolved full sociality, as other ecologically dominant fully social insects (e.g. ants, social bees and social wasps) have evolved from solitary predatory wasps.     

Evolution of Collective Behaviors for a Real Swarm of Aquatic Surface Robots

Swarm robotics is a promising approach for the coordination of large numbers of robots. While previous studies have shown that evolutionary robotics techniques can be applied to obtain robust and efficient self-organized behaviors for robot swarms, most studies have been conducted in simulation, and the few that have been conducted on real robots have been confined to laboratory environments. In this paper, we demonstrate for the first time a swarm robotics system with evolved control successfully operating in a real and uncontrolled environment. We evolve neural network-based controllers in simulation for canonical swarm robotics tasks, namely homing, dispersion, clustering, and monitoring. We then assess the performance of the controllers on a real swarm of up to ten aquatic surface robots. Our results show that the evolved controllers transfer successfully to real robots and achieve a performance similar to the performance obtained in simulation. We validate that the evolved controllers display key properties of swarm intelligence-based control, namely scalability, flexibility, and robustness on the real swarm. We conclude with a proof-of-concept experiment in which the swarm performs a complete environmental monitoring task by combining multiple evolved controllers.     

Paternal signature in kin recognition cues of a social insect: concealed in juveniles,revealed in adults

Kin recognition is a key mechanism to direct social behaviours towards related individuals or avoid inbreeding depression. In insects, recognition is generally mediated by cuticular hydrocarbon (CHC) compounds, which are partly inherited from parents. However, in social insects, potential nepotistic conflicts between group members from different patrilines are predicted to select against the expression of patriline-specific signatures in CHC profiles. Whereas this key prediction in the evolution of insect signalling received empirical support in eusocial insects, it remains unclear whether it can be generalized beyond eusociality to less-derived forms of social life. Here, we addressed this issue by manipulating the number of fathers siring clutches tended by females of the European earwig, Forficula auricularia, analysing the CHC profiles of the resulting juvenile and adult offspring, and using discriminant analysis to estimate the information content of CHC with respect to the maternal and paternal origin of individuals. As predicted, if paternally inherited cues are concealed during family life, increases in mating number had no effect on information content of CHC profiles among earwig juveniles, but significantly decreased the one among adult offspring. We suggest that age-dependent expression of patriline-specific cues evolved to limit the risks of nepotism as family-living juveniles and favour sibling-mating avoidance as group-living adults. These results highlight the role of parental care and social life in the evolution of chemical communication and recognition cues.     

Effects of aphids on foliar foraging by Argentine ants and the resulting effects on other arthropods

Abstract.  1. Although interactions between ants and honeydew-producing insects have received considerable study, relatively little is known about how these interactions alter the behaviour of ants in ways that affect other arthropods. In this study, field and greenhouse experiments were performed that examined how the presence of aphids ( Aphis fabae solanella ) on Solanum nigrum influenced the foraging behaviour of Argentine ants ( Linepithema humile ) and, in turn, modified the extent to which ants deter larval lacewings ( Chrysoperla rufilabris ), which are known aphid predators.
2. A field experiment demonstrated that the level of foliar foraging by ants increased linearly with aphid abundance, whereas no relationship existed between the level of ground foraging by ants and aphid abundance.
3. In the greenhouse, as in the field, foliar foraging by ants greatly increased when aphids were present. Higher levels of foliar foraging led to a twofold increase in the likelihood that ants contacted aphid predators. As a result of these increased encounters with ants, lacewing larvae were twice as likely to be removed from plants with aphids compared with plants without aphids. Once contact was made, however, the behaviour of ants towards lacewing larvae appeared similar between the two experimental groups.
4. Argentine ants drive away or prey upon a diversity of arthropod predators and parasitoids, but they also exhibit aggression towards certain herbivores. Future work should attempt to quantify how the ecological effects that result from interactions between honeydew-producing insects and invasive ants, such as L. humile , differ from those that result from interactions between honeydew-producing insects and native ants.     

Forage collection,substrate preparation,and diet composition in fungus‐growing ants

1. Variation and control of nutritional input is an important selective force in the evolution of mutualistic interactions and may significantly affect coevolutionary modifications in partner species. 2. The attine fungus‐growing ants are a tribe of more than 230 described species (12 genera) that use a variety of different substrates to manure the symbiotic fungus they cultivate inside the nest. Common ‘wisdom’ is that the conspicuous leaf‐cutting ants primarily use freshly cut plant material, whereas most of the other attine species use dry and partly degraded plant material such as leaf litter and caterpillar frass, but systematic comparative studies of actual resource acquisition across the attine ants have not been done. 3. Here we review 179 literature records of diet composition across the extant genera of fungus‐growing ants. The records confirm the dependence of leaf‐cutting ants on fresh vegetation but find that flowers, dry plant debris, seeds (husks), and insect frass are used by all genera, whereas other substrates such as nectar and insect carcasses are only used by some. 4. Diet composition was significantly correlated with ant substrate preparation behaviours before adding forage to the fungus garden, indicating that diet composition and farming practices have co‐evolved. Neither diet nor preparation behaviours changed when a clade within the paleoattine genus Apterostigma shifted from rearing leucocoprinous fungi to cultivating pterulaceous fungi, but the evolutionary derived transition to yeast growing in the Cyphomyrmex rimosus group, which relies almost exclusively on nectar and insect frass, was associated with specific changes in diet composition. 5. The co‐evolutionary transitions in diet composition across the genera of attine ants indicate that fungus‐farming insect societies have the possibility to obtain more optimal fungal crops via artificial selection, analogous to documented practice in human subsistence farming.     

Adaptive social immunity in leaf-cutting ants

Social insects have evolved a suite of sophisticated defences against parasites. In addition to the individual physiological immune response, social insects also express ‘social immunity’ consisting of group-level defences and behaviours that include allogrooming. Here we investigate whether the social immune response of the leaf-cutting ant Acromyrmex echinatior reacts adaptively to the virulent fungal parasite, Metarhizium anisopliae. We ‘immunized’ mini-nests of the ants by exposing them twice to the parasite and then compared their social immune response with that of naive mini-nests that had not been experimentally exposed to the parasite. Ants allogroomed individuals exposed to the parasite, doing this both for those freshly treated with the parasite, which were infectious but not yet infected, and for those treated 2 days previously, which were already infected but no longer infectious. We found that ants exposed to the parasite received more allogrooming in immunized mini-nests than in naive mini-nests. This increased the survival of the freshly treated ants, but not those that were already infected. The results thus indicate that the social immune response of this leaf-cutting ant is adaptive, with the group exhibiting a greater and more effective response to a parasite that it has previously been exposed to.     

Weeding and grooming of pathogens in agriculture by ants

The ancient mutualism between fungus-growing ants and the fungi they cultivate for food is a textbook example of symbiosis. Fungus-growing ants' ability to cultivate fungi depends on protection of the garden from the aggressive microbes associated with the substrate added to the garden as well as from the specialized virulent garden parasite Escovopsis. We examined ants' ability to remove alien microbes physically by infecting Atta colombica gardens with the generalist pathogen Trichoderma viride and the specialist pathogen Escovopsis. The ants sanitized the garden using two main behaviours: grooming of alien spores from the garden (fungus grooming) and removal of infected garden substrate (weeding). Unlike previously described hygienic behaviours (e.g. licking and self-grooming), fungus-grooming and garden-removal behaviours are specific responses to the presence of fungal pathogens. In the presence of pathogens, they are the primary activities performed by workers, but they are uncommon in uninfected gardens. In fact, workers rapidly eliminate Trichoderma from their gardens by fungus grooming and weeding, suggesting that these behaviours are the primary method of garden defence against generalist pathogens. The same sanitary behaviours were performed in response to the presence of the specialist pathogen Escovopsis. However, the intensity and duration of these behaviours were much greater in this treatment. Despite the increased effort, the ants were unable to eliminate Escovopsis from their gardens, suggesting that this specialized pathogen has evolved counter-adaptations in order to overcome the sanitary defences of the ants.     

High mating frequency and variation with lineage ratio in dependent-lineage harvester ants

Explaining the evolution of multiple mating is a challenge because of the associated costs. For social insects, mating frequency may have fitness consequences due to effects on social interactions or genetic diversity within colonies. Here, we investigated the evolution of mating frequency in a social insect species with a unique genetic system that requires multiple mating. In certain populations of Pogonomyrmex harvester ants, there are two interbreeding yet genetically distinct mitochondrial lineages. Queens must mate with males of the opposite lineage to produce workers and with males of the same lineage to produce reproductive females. We expected queens of the dependent-lineage system to exhibit high mating frequencies relative to other social insects. Furthermore, we expected queens from populations of highly asymmetric lineage ratios to exhibit even higher mating frequencies, to adequately sample the population and successfully mate with males of the less common lineage. To test these predictions, we estimated the mating frequency of 16 P. barbatus queens, and compared these mating frequencies between two populations, one with relatively equal lineage ratio (60:40) and a second with a highly asymmetrical lineage ratio (96:4). Overall, observed mating frequency exceeded 10, which is high in comparison to other social insects, and our estimates of effective mating frequency were among the highest of Pogonomyrmex species. Mating frequency at the site with the asymmetrical lineage ratio was also significantly higher than the site with the more even ratio. Our results suggest that obligate multiple mating as well as lineage ratio contribute to the evolution of high mating frequency in dependent-lineage populations.     

Les symbioses entre plantes et fourmis arboricoles

In communities of tree-nesting ants in tropical rain forests, energy-rich resources from plants enable ants to achieve high densities and permit the evolution of energy-intensive strategies of prédation. Ants are more abundant, and can maintain populations of phytophagous insects at lower densities than if ant populations were limited simply by insect prey. A large proportion of woody plants in tropical forests are involved in such loose protection mutualisms. Many epiphytes are also involved in loose associations with ants, in which benefits to plants may include nutrition and seed dispersal as well as protection. From such opportunistic interactions numerous symbiotic mutualisms have evolved, in which ants and plants are more intimately and often more specifically associated. The diversity of these symbiotic interactions between ants and plants make these associations good models for examining many general questions in ecology. Plants that have evolved specialised structures (‘ant-domatia’) to house ants are termed ‘myrmécophytes’ or ‘ant-plants’, and their specialised associates are termed ‘plant-ants’. In these symbioses, plants and ants have coevolved. Ant colonies that provide increased benefits to the host plant enhance its growth and survival, thus receiving more benefits from it, and vice versa. Selection favours mutualistic traits, and interests of the two partners tend to converge. However, because these associations are horizontally transmitted, neither partner obtains benefits from reproduction of the other. Because reproduction draws away resources from growth (from which the partner benefits), it introduces conflicts of interest between ants and plants, and several examples show the importance of such conflicts in the dynamics of coevolution. Antplant coevolution has produced parasites as well as mutualists. Much is still unknown on the evolutionary ecology of these symbioses. Mechanisms of interaction at the chemical level (chemical ecology) are little explored. The functioning of ant-plant associations at the level of populations and communities is poorly understood, and information in this domain is crucial for the conservation of these intricate symbioses in forests increasingly subjected to disturbance and fragmentation.     

Social Insect Pheromones: Their Chemistry and Function

Exocrine secretions of social insects are often characterizedby extraordinarily complex mixtures of natural products. Thus,chemical communication in social insects must be interpretedin terms of signals generated by multicomponent systems, theindividual constituents of which can affect the informationalcontent of the message. Alarm pheromones have been identified chiefly in three subfamiliesof ants and their distribution appears to be chemosystematicallysignificant. Myrmicine genera emphasize 3-alkanones as alarmreleasers, whereas methyl ketones, primarily of terpenoidalorigin, are widely utilized as alarm pheromones in the subfamilyDolichoderinae. Formicine species may employ formic acidas analarm pheromone in addition to the compounds produced in themandibular and Dufour's glands. The mandibular gland pheromonesare chiefly acyclic monoterpene aldehydes (e.g., citronellal)which are relatively low boiling compounds. Higher boiling n-alkanesare produced in the Dufour's glands and may serve as more persistentreleasers of alarm behavior. Alarm pheromones as well as thecaste-specific pheromones of male bees and ants, probably alsoserve as defensive products. In many cases it is likely thatpheromones were originally utilized as defensive compounds andtheir communicative function is a secondary development.     

Cryptic termites avoid predatory ants by eavesdropping on vibrational cues from their footsteps

Eavesdropping has evolved in many predator–prey relationships. Communication signals of social species may be particularly vulnerable to eavesdropping, such as pheromones produced by ants, which are predators of termites. Termites communicate mostly by way of substrate‐borne vibrations, which suggest they may be able to eavesdrop, using two possible mechanisms: ant chemicals or ant vibrations. We observed termites foraging within millimetres of ants in the field, suggesting the evolution of specialised detection behaviours. We found the termite Coptotermes acinaciformis detected their major predator, the ant Iridomyrmex purpureus, through thin wood using only vibrational cues from walking, and not chemical signals. Comparison of 16 termite and ant species found the ants‐walking signals were up to 100 times higher than those of termites. Eavesdropping on passive walking signals explains the predator detection and foraging behaviours in this ancient relationship, which may be applicable to many other predator–prey relationships.     

Interactions between ants and aphidophagous and coccidophagous ladybirds

Aphidophagous and coccidophagous coccinellids come into conflict with homopteran-tending ants for access to food. Antagonistic interactions between coccinellids and ants may be competitive or non-competitive. Competitive interactions occur when coccinellids attack aphids or coccids that are being tended by ants for honeydew. Non-competitive interactions include all interactions away from ant-tended homopteran colonies. We here review observations and studies of such interactions. We note that most competitive interactions occur at times when untended aphids/coccids are scarce. We describe the chemical and physical defences that coccinellids use against ant aggression and consider whether these have evolved as general anti-predator deterrents or specifically in response to ants. Myrmecophilous coccinellids are then considered, with particular focus on the two most studied species, Coccinella magnifica and Platynaspis luteorubra. We note that the myrmecophily of the two species has the same adaptive rationale—to enable the ladybirds to prey on ant-tended aphids at times of aphid scarcity—but that it is based on different traits to facilitate life with ants. Finally, we consider the role of ants in the evolution of habitat specialisation in some coccinellids.     

Ecological interactions among insect herbivores,ants and the host plant Baccharis dracunculifolia in a Brazilian mountain ecosystem

Insect–plant interactions occur in several ways and have considerable environmental and ecological importance. Many feeding strategies have evolved among herbivorous insects, with host–herbivore systems likely being influenced by trophobionts with ants. We investigated how these interactions vary across elevation gradients by evaluating the structure of the herbivorous insect community and ants associated with Baccharis dracunculifolia at three distinct elevations (800, 1100, and 1400 m a.s.l.) on a mountain in southeastern Brazil. Moreover, we evaluated the diversity and specialisation of interactions between herbivores and host plants along the elevational gradient. We sampled herbivores and ants on 60 plants at each elevation (totalling 180 plant individuals). Herbivore species composition differed among elevations, as did interaction diversity and specialisation. Richness and abundance of chewing insects increased with elevation, while β‐diversity among patches of the host plant was higher at the lowest elevation, probably due to the patchy occurrence of B. dracunculifolia. Richness and abundance of sap‐sucking insects were higher at the intermediate elevation, possibly due to local environmental conditions. We observed a positive relationship between ant and herbivore trophobiont richness on B. dracunculifolia. We found that interactions were more specialised and less diverse at higher elevations compared to the lowest elevation. Changes in vegetation and environmental variables shaped species distributions and their ecological interactions along the elevation gradient. Our study demonstrates that increased elevation changes the structure and patterns of interactions of the herbivore insect guilds associated with the host plant B. dracunculifolia. Ant effects depend on the context, the environment, and the species of ants involved, and are essential for the presence of insect trophobionts.     

Robot control and the evolution of modular neurodynamics

Summary A modular approach to neural behavior control of autonomous robots is presented. It is based on the assumption that complex internal dynamics of recurrent neural networks can efficiently solve complex behavior tasks. For the development of appropriate neural control structures an evolutionary algorithm is introduced, which is able to generate neuromodules with specific functional properties, as well as the connectivity structure for a modular synthesis of such modules. This so called ENS ³-algorithm does not use genetic coding. It is primarily designed to develop size and connectivity structure of neuro-controllers. But at the same time it optimizes also parameters of individual networks like synaptic weights and bias terms. For demonstration, evolved networks for the control of miniature Khepera robots are presented. The aim is to develop robust controllers in the sense that neuro-controllers evolved in a simulator show comparably good behavior when loaded to a real robot acting in a physical environment. Discussed examples of such controllers generate obstacle avoidance and phototropic behaviors in non-trivial environments.