Does ant predation favour leaf beetle specialization on toxic host plants?

Generalist predators are frequently seen as evolutionary forces that narrow the host range in herbivorous insects. Predators may favour specialization of herbivores on host plants containing toxic chemicals (which can be used by herbivores for their own defence) if host plant‐derived defences provide better protection from enemies than do autogenously produced defences. We compared the effectiveness of these two defensive strategies in the larvae of six species of leaf beetle (Chrysomelidae) against wood ants (Formica rufa group) in field experiments. Ants were more strongly repelled by larvae with host plant‐derived, salicylaldehyde‐containing secretions than by larvae with various autogenous secretions, but collectively foraging ants ultimately overcame any type of chemical defence by social interactions, chemical signalling, and olfactory learning. As a result, ants killed all larvae of Chrysomela lapponica defended by salicylaldehyde‐containing secretions within 2 days of their introduction to willows within 15 m of ant nests. We conclude that in the field neither type of chemical defence provides complete protection against wood ants in the vicinity of their nests, and that evolutionary shifts from autogenous production of secretion to sequestration of plant allelochemicals in leaf beetles may be favoured mostly at low ant densities on the periphery of ant foraging areas.     

Strategies of the beetle Oochrotus unicolor (Tenebrionidae) thriving in the waste dumps of seed-harvesting Messor ants (Formicidae)

1. A diverse group of arthropods have adapted to the niches found inside the nests of social insects. Studies mostly focused on very specialised parasites residing in the brood chambers. However, the biology and strategies of symbionts occupying other niches, such as waste dumps, are underexplored. 2. Using a series of complementary experiments, this study demonstrated that the Mediterranean beetle Oochrotus unicolor has adapted to the waste dump niche found in the nests of Messor harvester ants. 3. Laboratory experiments confirmed field observations that the beetle preferentially resided in the refuse pits. Next, it was shown that the beetles readily consumed seeds and flour, whereas other food sources were poorly accepted and ant brood was never even eaten. The beetles did not elicit a strong aggression response in Messor ants, and they could tolerate very high densities of workers without clear costs. The beetles modestly mimicked the nest recognition cues of their Messor host. This imperfect mimicry could promote the adoption of the beetle in the ant colony, in concert with mechanical defence generated by its tank-like body. Isolation of the beetle from its host did not significantly affect the beetle's chemical cuticular profile nor did it provoke elevated ant aggression, indicating that the beetle does not acquire the chemicals passively from its host. 4. This paper discusses the fact that waste dumps in social insect nests are hotspots for arthropod symbionts. It shows that symbionts in this niche may employ behavioural, trophic and chemical strategies that are different from those found in other niches of social insect nests.     

Cascading trait‐mediated interactions induced by ant pheromones

Trait‐mediated indirect interactions (TMII) can be as important as density‐mediated indirect interactions. Here, we provide evidence for a novel trait‐mediated cascade (where one TMII affects another TMII) and demonstrate that the mechanism consists of a predator eavesdropping on chemical signaling. Ants protect scale insects from predation by adult coccinellid beetles – the first TMII. However, parasitic phorid flies reduce ant foraging activity by 50% – the second TMII, providing a window of opportunity for female beetles to oviposit in high‐quality microsites. Beetle larvae are protected from ant predation and benefit from living in patches with high scale densities. We demonstrate that female beetles can detect pheromones released by the ant when attacked by phorids, and that only females, and especially gravid females, are attracted to the ant pheromone. As ants reduce their movement when under attack by phorids, we conclude that phorids facilitate beetle oviposition, thus producing the TMII cascade.     

The contribution of cacao agroforests to the conservation of lower canopy ant and beetle diversity in Indonesia

The ongoing destruction of tropical rainforests has increased the interest in the potential value of tropical agroforests for the conservation of biodiversity. Traditional, shaded agroforests may support high levels of biodiversity, for some groups even approaching that of undisturbed tropical forests. However, it is unclear to what extent forest fauna is represented in this diversity and how management affects forest fauna in agroforests. We studied lower canopy ant and beetle fauna in cacao agroforests and forests in Central Sulawesi, Indonesia, a region dominated by cacao agroforestry. We compared ant and beetle species richness and composition in forests and cacao agroforests and studied the impact of two aspects of management intensification (the decrease in shade tree diversity and in shade canopy cover) on ant and beetle diversity. The agroforests had three types of shade that represented a decrease in tree diversity (high, intermediate and low diversity). Species richness of ants and beetles in the canopies of the cacao trees was similar to that found in lower canopy forest trees. However, the composition of ant and beetle communities differed greatly between the agroforest and forest sites. Forest beetles suffered profoundly from the conversion to agroforests: only 12.5% of the beetle species recorded in the forest sites were also found in the agroforests and those species made up only 5% of all beetles collected from cacao. In contrast, forest ants were well represented in agroforests, with 75% of all species encountered in the forest sites also occurring on cacao. The reduction of shade tree diversity had no negative effect on ants and beetles on cacao trees. Beetle abundances and non-forest ant species richness even increased with decreasing shade tree diversity. Thinning of the shade canopy was related to a decrease in richness of forest ant species on cacao trees but not of beetles. The contrasting responses of ants and beetles to shade tree management emphasize that conservation plans that focus on one taxonomic group may not work for others. Overall ant and beetle diversity can remain high in shaded agroforests but the conservation of forest ants and beetles in particular depends primarily on the protection of natural forests, which for forest ants can be complemented by the conservation of adjacent shaded cacao agroforests.     

Landscape effects of transgenic cotton on non-target ants and beetles

Transgenic crops producing toxins from Bacillus thuringiensis (Bt) can be planted in the same field for many years, and many insects exploiting such crops must disperse to other habitats to persist. Accordingly, effects of transgenic crop farming could accumulate through time and affect insect populations across agricultural landscapes. We monitored the population density of seven ant genera and beetle families and of rare ants and beetles in 84 non-cultivated sites abutting agricultural fields in Central Arizona. We assessed the short-term (during planting year) and long-term (over 5–6 years) landscape effect of farming Cry1Ac cotton on ant and beetle density in non-cultivated sites, in addition to several local and regional variables. Landscape variables (e.g., sequence of crops planted in neighbouring fields, crop diversity, and abundance) were more frequently associated with insect density than local variables (e.g., plant productivity and diversity in non-cultivated sites). In the short-term, use of Bt relative to non-Bt cotton in neighbouring fields was positively associated with density of one ant and two beetle groups in non-cultivated sites. However, acreage of Bt cotton located within 1 km from non-cultivated sites had more negative effects than acreage of non-Bt cotton on density of one ant and one beetle group. In the long-term, the proportion of years that Bt cotton was planted in neighbouring fields was positively associated with ant density but not beetle density. Results suggest that the farming of Bt cotton in neighbouring fields frequently resulted in positive short- and long-term landscape effects on ants and beetles in non-cultivated sites, while Bt cotton planted farther away had less frequent negative short-term impacts.     

The rove beetle Drusilla sparsa (Coleoptera: Staphylinidae) is a myrmecophilous species associated with a myrmicine ant,Crematogaster osakensis (Hymenoptera: Formicidae)

The rove beetle genus Drusilla includes some myrmecophilous species. The Japanese species Drusilla sparsa (Sharp, 1874) has been regarded as a non‐myrmecophilous beetle. In Kagawa Prefecture, Shikoku Island, western Japan, however, we often observed that D. sparsa adults were walking in the vicinity of foraging workers of the myrmicine ant Crematogaster osakensis Forel, 1990. The body color of the beetle is similar to C. osakensis as in other myrmecophilous beetles found near the trails of the host ants. To examine whether D. sparsa is myrmecophilous, we investigated the distribution of D. sparsa and C. osakensis in the field, as well as their behavior including prey preference of the beetle in the laboratory. Drusilla sparsa beetles were collected only in sites where C. osakensis ants occurred. When the beetles encountered the ant workers, they bent the abdominal tip toward the ants. The ants licked the abdominal tip, and then the beetles usually walked away. Such behavioral reaction of the ants was not observed when the beetles encountered workers of the formicine ant Nylanderia flavipes (Smith, 1874) that continuously attacked the beetles. Drusilla sparsa preferred to feed on dead workers of C. osakensis even when other ants were available as food, indicating that D. sparsa is a myrmecophilous species associated with C. osakensis. Crematogaster osakensis was frequently found in the stomach in the ant predator, the Japanese treefrog Hyla japonica Günther, 1859. Thus, the significance of body color similarity between the host ants and beetles is not a case of Batesian mimicry.     

Behavioral mechanisms of spatial competition between red wood ants (Formica aquilonia) and ground beetles (Carabidae)

Behavioral aspects of spatial competition between red wood ants (Formica aquilonia) and six mass species of Carabidae were studied in field and laboratory experiments. We showed that red wood ants essentially influence spatial distribution of ground beetles on their common territories. Transplantation experiments suggest that in newly established ants' settlements stronger forms of interrelations arise than in old stable colony. To examine the ability of beetles to avoid collisions with ants we used two experimental techniques. In laboratory, we tested carabids ability to avoid a clash in a Y-shaped labyrinth containing an active tethered ant in one section. In field experiments we compared quantitative characteristics of movements (such as crookedness of individual trajectories, speed of movement, the time spent on stops) for beetles placed close to ants foraging routes and on ant-free plots. All beetles studied displayed a clear tendency to learn, that is, to modity their behavior in order to avoid collisions with ants. Species that exhibited best parameters of learning were closer to ants by their size and characteristic movement, namely, Pterostichus oblogopunctatus and P. magus. Beetles' stereotyped behavioral tactics can be considered universal for avoiding collisions with any subject (for instance, with an ant) of a certain size and speed of movements. A set of tactics in the labyrinth included: (1) attempts to round the ant; (2) turns away after touching the ant with antennae; (3) turns away without a contact; (4) avoidances of a dangerous section; (5) stops near the ant with the antennae hidden. Comparing pairwise difference between four species shows that beetles use species-specific preference for definite combinations of tactics. Effective learning allows carabids to penetrate into ant foraging territory and partly avoide interference competition. It seems that red wood ants are not inclined to learn to avoid collisions with competing carabid species. Instead, they recognize an "enemy's image" and selectively attack relatively small predatory carabids rather than herbivorous species. Experiments with dummy beetles suggest that ants react for several hierarchically organized key characteristics of competitors such as speed of movement, dark color, bilateral symmetry, protuberances (legs, antennae), and smell. Among "professional" groups in ant family, guards and hunters react to beetles aggressively, whereas aphid tenders ignore them. Sophisticated combination of flexible and innate behavioral patterns enables insects to share territories and interact in the mode of relatively mild, spatial, competition instead of predation. Eliciting sets of hierarchically organized features of competitors that govern adjustment of spatial distribution in insects' community enable us to integrate ideas of classic and cognitive ethology.     

Wide corridors with much environmental heterogeneity best conserve high dung beetle and ant diversity

Landscape ecological networks (ENs) consist of landscape-scale conservation corridors that connect areas of high natural value within a production mosaic with protected areas (PAs). In South Africa, ENs have been implemented on a large spatial scale to offset the negative impacts of plantation forestry on indigenous grasslands. We focus on corridor width as a factor for conserving dung beetle and ant diversity within an EN. We also investigate the importance of natural environmental heterogeneity (elevation, vegetation type) and habitat quality (soil hardness, invasive alien plant density). We sampled dung beetles and ants in 30 corridors of different sizes, and at ten sites in a nearby PA. In addition, we also analysed dung beetles according to their feeding guild. Tunnelling dung beetle species richness increased with corridor width. Rolling dung beetle species richness was higher in the PA than in the corridors of the EN. The dung beetle assemblage within the EN differed from that within the PA. Corridors of various widths differed in ant composition but not in species richness. Furthermore, the PA and the EN differed in environmental variables, which contributed to differences in dung beetle species richness and assemblage composition. Within the EN, environmental heterogeneity across the landscape was more important than corridor width for driving species diversity of both dung beetles and ants. When planning future ENs, wide corridors (>280 m) that encompass as much natural heterogeneity across the landscape as possible will best conserve the range of local insect species.     

Cryptic diversity,high host specificity and reproductive synchronization in army ant‐associated Vatesus beetles

Army ants and their arthropod symbionts represent one of the most species‐rich animal associations on Earth, and constitute a fascinating example of diverse host–symbiont interaction networks. However, despite decades of research, our knowledge of army ant symbionts remains fragmentary due to taxonomic ambiguity and the inability to study army ants in the laboratory. Here, we present an integrative approach that allows us to reliably determine species boundaries, assess biodiversity, match different developmental stages and sexes, and to study the life cycles of army ant symbionts. This approach is based on a combination of community sampling, DNA barcoding, morphology and physiology. As a test case, we applied this approach to the staphylinid beetle genus Vatesus and its different Eciton army ant host species at La Selva Biological Station, Costa Rica. DNA barcoding led to the discovery of cryptic biodiversity and, in combination with extensive community sampling, revealed strict host partitioning with no overlap in host range. Using DNA barcoding, we were also able to match the larval stages of all focal Vatesus species. In combination with studies of female reproductive physiology, this allowed us to reconstruct almost the complete life cycles of the different beetle species. We show that Vatesus beetles are highly adapted to the symbiosis with army ants, in that their reproduction and larval development are synchronized with the stereotypical reproductive and behavioural cycles of their host colonies. Our approach can now be used to study army ant‐symbiont communities more broadly, and to obtain novel insights into co‐evolutionary and ecological dynamics in species‐rich host–symbiont systems.     

Discovery of defense- and neuropeptides in social ants by genome-mining

Natural peptides of great number and diversity occur in all organisms, but analyzing their peptidome is often difficult. With natural product drug discovery in mind, we devised a genome-mining approach to identify defense- and neuropeptides in the genomes of social ants from Atta cephalotes (leaf-cutter ant), Camponotus floridanus (carpenter ant) and Harpegnathos saltator (basal genus). Numerous peptide-encoding genes of defense peptides, in particular defensins, and neuropeptides or regulatory peptide hormones, such as allatostatins and tachykinins, were identified and analyzed. Most interestingly we annotated genes that encode oxytocin/vasopressin-related peptides (inotocins) and their putative receptors. This is the first piece of evidence for the existence of this nonapeptide hormone system in ants (Formicidae) and supports recent findings in Tribolium castaneum (red flour beetle) and Nasonia vitripennis (parasitoid wasp), and therefore its confinement to some basal holometabolous insects. By contrast, the absence of the inotocin hormone system in Apis mellifera (honeybee), another closely-related member of the eusocial Hymenoptera clade, establishes the basis for future studies on the molecular evolution and physiological function of oxytocin/vasopressin-related peptides (vasotocin nonapeptide family) and their receptors in social insects. Particularly the identification of ant inotocin and defensin peptide sequences will provide a basis for future pharmacological characterization in the quest for potent and selective lead compounds of therapeutic value.     

Task-specific expression of the foraging gene in harvester ants

In social insects, groups of workers perform various tasks such as brood care and foraging. Transitions in workers from one task to another are important in the organization and ecological success of colonies. Regulation of genetic pathways can lead to plasticity in social insect task behaviour. The colony organization of advanced eusocial insects evolved independently in ants, bees, and wasps and it is not known whether the genetic mechanisms that influence behavioural plasticity are conserved across species. Here we show that a gene associated with foraging behaviour is conserved across social insect species, but the expression patterns of this gene are not. We cloned the red harvester ant (Pogonomyrmex barbatus) ortholog (Pbfor) to foraging, one of few genes implicated in social organization, and found that foraging behaviour in harvester ants is associated with the expression of this gene; young (callow) worker brains have significantly higher levels of Pbfor mRNA than foragers. Levels of Pbfor mRNA in other worker task groups vary among harvester ant colonies. However, foragers always have the lowest expression levels compared to other task groups. The association between foraging behaviour and the foraging gene is conserved across social insects but ants and bees have an inverse relationship between foraging expression and behaviour.     

Ant diversity as a direct and indirect driver of pselaphine rove beetle (Coleoptera: Staphylinidae) functional diversity in tropical rainforests,Sabah, Malaysian Borneo

Pselaphinae is a species‐rich beetle subfamily found globally, with many exhibiting myrmecophily—a symbiotic association with ants. Pselaphine–ant associations vary from facultative to obligate, but direct behavioral observations still remain scarce. Pselaphines are speciose and ecologically abundant within tropical leaf litter invertebrate communities where ants dominate, implying a potentially important ecological role that may be affected by habitat disturbances that impact ants. In this study, we measured and analyzed putative functional traits of leaf litter pselaphines associated with myrmecophily through morphometric analysis. We calculated “myrmecophile functional diversity” of pselaphines at different sites and examined this measure's relationship with ant abundance, in both old growth and logged rainforest sites in Sabah, Borneo. We show that myrmecophile functional diversity of pselaphine beetles increases as ant abundance increases. Old growth rainforest sites support a high abundance of ants, which is associated with a high abundance of probable myrmecophilous pselaphines. These results suggest a potential link between adult morphological characters and the functional role these beetles play in rainforest litter as ecological interaction partners with ants.     

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.     

Interactional behaviors of the parasitic beetle Paussus favieri with its ant host Pheidole pallidula: the mimetic role of the acoustical signals

The social parasitic beetle Puussus favieri(Coleoptera,Carabidae,Paussini)performs different types of stridulations,which sclectively mimic those emitted by dif-ferent ant castes of its host Pheidole pallidula(Hymenoptera,Formicidae,Myrmicinae).However,the significance of this acoustical mimicry for the success of the parasitic strat-egy and the behaviors elicited in the host ants by stridulations was unknown.We reared Paussus favieri in Pheidole pallidula colonies and filmed their interacting behaviors.We analyzed in slow motion the behavior of ants near a stridulating beetle.We analyzed sep-arately trains of pulse(Pa+Pb,produced by repeated rubbings)and single pulse(Pc,produced by a single rubbing)of stridulations,clearly recognizable from the shaking up and down of the beetle hind legs.and associated them with differcent ant responscs.The full repertoire of sounds produced by P:favieri elicited benevolent responses both in workers and soldiers.We found that different signals elicit different(sometimes multiplc)bchaviors in ants,with different frequency in the two ant castes.However,Pc(alone or in conjunction with other types of pulses)appears to be the type of acoustic signal mostly responsible for all recorded behaviors.These results indicate that the acoustic channel plays a pivotal role in the host-parasite interaction.Finding that a parasite uses the acoustical channel so intensively,and in such a complicated way to trigger ant bchaviors,indicates that acoustic signals may be more important in ant societies than commonly recognized.     

The indirect consequences of a mutualism: comparing positive and negative components of the net interaction between honeydew-tending ants and host plants

1. In ecological webs, net indirect interactions between species are composed of interactions that vary in sign and magnitude. Most studies have focused on negative component interactions (e.g. predation, herbivory) without considering the relative importance of positive interactions (e.g. mutualism, facilitation) for determining net indirect effects. 2. In plant/arthropod communities, ants have multiple top-down effects via mutualisms with honeydew-producing herbivores and harassment of and predation on other herbivores; these ant effects provide opportunities for testing the relative importance of positive and negative interspecific interactions. We manipulated the presence of ants, honeydew-producing membracids and leaf-chewing beetles on perennial host plants in field experiments in Colorado to quantify the relative strength of these different types of interactions and their impact on the ant's net indirect effect on plants. 3. In 2007, we demonstrated that ants simultaneously had a positive effect on membracids and a negative effect on beetles, resulting in less beetle damage on plants hosting the mutualism. 4. In 2008, we used structural equation modelling to describe interaction strengths through the entire insect herbivore community on plants with and without ants. The ant's mutualism with membracids was the sole strong interaction contributing to the net indirect effect of ants on plants. Predation, herbivory and facilitation were weak, and the net effect of ants reduced plant reproduction. This net indirect effect was also partially because of behavioural changes of herbivores in the presence of ants. An additional membracid manipulation showed that the membracid's effect on ant activity was largely responsible for the ant's net effect on plants; ant workers were nearly ten times as abundant on plants with mutualists, and effects on other herbivores were similar to those in the ant manipulation experiment. 5. These results demonstrate that mutualisms can be strong relative to negative direct interspecific interactions and that positive interactions deserve attention as important components of ecological webs.     

An experimental study of behavioral mechanisms of relationships between red wood ants and ground beetles

Behavioral mechanisms regulating the competitive relations between the red wood ant Formica aquilonia and the ground beetles Carabus regalis, Pterostichus melanarius, P. magus, P. oblongopunctatus, Harpalus smaragdinus, and Amara nitida were studied by artificially initiated collisions between living insects as well as with the use of imitation models. Members of different functional groups within an ant family (aphid-milkers, hunters, and guards) behaved differently towards beetles. Active ants were shown to respond selectively to different features of the possible competitors, such as coloration, the presence of “appendages” (legs, antennae), body symmetry, rate of movement, and scent. Field and laboratory experiments demonstrated the ability of beetles to avoid collisions with active ants. The scent of anthill material attracted ground beetles, which consumed dead ants. The gained individual experience may allow the beetles to use supplementary forage resources in the territories controlled by ants. The flexible tactical patterns facilitate spatial segregation of ground beetles and ants in the same territory and result in a more complete utilization of food resources.     

Lethal and sublethal effects of thiacloprid on non-target carpenter ant,Camponotus japonicus Mayr (Hymenoptera: Formicidae)

To control population of Monochamus beetles that transmit pine wood nematode, Bursaphelenchus xylophilus, a variety of insecticides have been applied to forest ecosystems in Korea. Non-target predatory insects can be directly or indirectly exposed to insecticides. We evaluated potential lethal and sublethal effects of thiacloprid on survival and behavior of carpenter ants, Camponotus japonicus Mayr. Field-collected ants were directly exposed to several food items such as thiacloprid-exposed Monochamus beetles, 10% sugar water with thiacloprid concentrations, and 10% sugar water at group and individual levels. In experiment for groups of individuals, dead beetle bodies generally had possible adverse effects on ants through dietary exposure, because two forager ants were dead or paralysis after they were exposed to thiacloprid-exposed Monochamus beetles. At individual level, dietary exposure to thiacloprid at concentrations of 10 and 50?mg/L was lethal to ants, causing paralysis and impaired walking. Mortality of ant workers was higher in direct or indirect exposure treatments than that in the control. Some ants exposed to thiacloprid showed abnormal behavior within a few days, especially at thiacloprid concentrations of 10 and 50?mg/L. However, some of them sometimes recovered from the abnormal behavior in a day. In consequence, application of thiacloprid in pine forests may disrupt species interaction and foraging behavior of ants, but the effect of thiacloprid through trophallaxis behavior should be further assessed using more extensive colonies composed of queen, workers, brood, and so on.     

Do pygidial secretions of dung beetles have the potential to repel urban pest ants?

Various organisms emit malodorous secretions against competitors, and the potential use of these secretions in pest management should be investigated. For example, some ant species feed on similar resources as dung beetles, which might have led to counter chemical defences in dung beetles. We tested whether pygidial secretions of the dung beetle Canthon smaragdulus (Fabricius) (Coleoptera: Scarabaeidae, Scarabaeinae) alter the locomotor behaviour of the exotic urban pest ant Tapinoma melanocephalum (Fabricius) (Hymenoptera: Formicidae), specifically whether these secretions repel those ants. We also tested whether the disturbance in the locomotor behaviour of T. melanocephalum increases with the amount of pygidial secretion. We found that individual T. melanocephalum displayed changes in their locomotor behaviour when exposed to pygidial secretions of coupled dung beetles, single males, and single females. Additionally, the pygidial secretions from male and female dung beetles could repel ants. The change in the locomotor behaviour of T. melanocephalum increased with the amount of pygidial secretion. Our results suggest that the pygidial secretions of dung beetles have potential as a biological repellent of T. melanocephalum. Hence, pygidial secretions from dung beetles may be used in the future for the development of urban pest management strategies.     

Sick ants become unsociable

Parasites represent a severe threat to social insects, which form high-density colonies of related individuals, and selection should favour host traits that reduce infection risk. Here, using a carpenter ant (Camponotus aethiops) and a generalist insect pathogenic fungus (Metarhizium brunneum), we show that infected ants radically change their behaviour over time to reduce the risk of colony infection. Infected individuals (i) performed less social interactions than their uninfected counterparts, (ii) did not interact with brood anymore and (iii) spent most of their time outside the nest from day 3 post-infection until death. Furthermore, infected ants displayed an increased aggressiveness towards non-nestmates. Finally, infected ants did not alter their cuticular chemical profile, suggesting that infected individuals do not signal their physiological status to nestmates. Our results provide evidence for the evolution of unsociability following pathogen infection in a social animal and suggest an important role of inclusive fitness in driving such evolution.     

Social encapsulation of the small hive beetle (<Emphasis Type="Italic">Aethina tumida</Emphasis> Murray) by European honeybees (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

Summary European and African subspecies of honeybees (Apis mellifera L.) utilize social encapsulation to contain the small hive beetle (Aethina tumida Murray), a honeybee colony scavenger. Using social encapsulation, African honeybees successfully limit beetle reproduction that can devastate host colonies. In sharp contrast, European honeybees often fail to contain beetles, possibly because their social encapsulation skills may be less developed than those of African honeybees. In this study, we quantify beetle and European honeybee behaviours associated with social encapsulation, describe colony and time (morning and evening) differences in these behaviours (to identify possible circadian rhythms), and detail intra-colonial, encapsulated beetle distributions. The data help explain the susceptibility of European honeybees to depredation by small hive beetles. There were significant colony differences in a number of social encapsulation behaviours (the number of beetle prisons and beetles per prison, and the proportion of prison guard bees biting at encapsulated beetles) suggesting that successful encapsulation of beetles by European bees varies between colonies. We also found evidence for the existence of circadian rhythms in small hive beetles, as they were more active in the evening rather than morning. In response to increased beetle activity during the evening, there was an increase in the number of prison guard bees during evening. Additionally, the bees successfully kept most (~93%) beetles out of the combs at all times, suggesting that social encapsulation by European honeybees is sufficient to control small populations of beetles (as seen in this study) but may ultimately fail if beetle populations are high.Received 20 January 2003; revised 21 April 2003; accepted 29 April 2003.