When attempts at robbing prey turn fatal

Because group-hunting arboreal ants spread-eagle insect prey for a long time before retrieving them, these prey can be coveted by predatory flying insects. Yet, attempting to rob these prey is risky if the ant species is also an effective predator. Here, we show that trying to rob prey from Azteca andreae workers is a fatal error as 268 out of 276 potential cleptobionts (97.1?%) were captured in turn. The ant workers hunt in a group and use the “Velcro?” principle to cling firmly to the leaves of their host tree, permitting them to capture very large prey. Exceptions were one social wasp, plus some Trigona spp. workers and flies that landed directly on the prey and were able to take off immediately when attacked. We conclude that in this situation, previously captured prey attract potential cleptobionts that are captured in turn in most of the cases.     

Low chitinase activity in Acacia myrmecophytes: a potential trade-off between biotic and chemical defences?

 We determined chitinase activity in leaves of four myrmecophytic and four non-myrmecophytic leguminous species at the plants' natural growing sites in Mexico. Myrmecophytic plants (or 'ant plants') have obligate mutualisms with ants protecting them against herbivores and pathogenic fungi. Plant chitinases can be considered a reliable measure of plant resistance to pathogenic fungi. The myrmecophytic Acacia species, which were colonised by mutualistic ants, exhibited at least six-fold lower levels of chitinase activity compared with the non-myrmecophytic Acacia farnesiana and three other non-myrmecophytes. Though belonging to different phylogenetic groups, the myrmecophytic Acacia species formed one distinct group in the data set, which was clearly separated from the non-myrmecophytic species. These findings allowed for comparison between two recent hypotheses that attempt to explain low chitinase activity in ant plants. Most probably, chitinases are reduced in myrmecophytic plant species because these are effectively defended indirectly due to their symbiosis with mutualistic ants. Received: 16 August 2000 / Accepted in revised form: 28 October 2000     

Predation and aggressiveness in host plant protection: a generalization using ants from the genus Azteca

In studying the ant genus Azteca, a Neotropical group of arboreal species, we aimed to determine the extent to which the ants use predation and/or aggressiveness to protect their host plants from defoliating insects. We compared a territorially dominant, carton-nester, Azteca chartifex, and three plant-ant species. Azteca alfari and Azteca ovaticeps are associated with the myrmecophyte Cecropia (Cecropiaceae) and their colonies shelter in its hollow branches; whereas Azteca bequaerti is associated with Tococa guianensis (Melastomataceae) and its colonies shelter in leaf pouches situated at the base of the laminas. Whereas A. bequaerti workers react to the vibrations transmitted by the lamina when an alien insect lands on a leaf making it unnecessary for them to patrol their plant, the workers of the three other species rather discover prey by contact. The workers of all four species use a predatory behaviour involving spread-eagling alien insects after recruiting nestmates at short range, and, in some cases, at long range. Because A. alfari and A. ovaticeps discard part of the insects they kill, we deduced that the workers' predatory behaviour and territorial aggressiveness combine in the biotic defence of their host tree.     

A unique strategy of host colony exploitation in a parasitic ant: workers of<Emphasis Type="Italic"> Polyrhachis lama</Emphasis> rear their brood in neighbouring host nests

The formicine ant Polyrhachis lama is a social parasite, exploiting its ponerine host ant species Diacamma sp. In most social parasitic associations, the parasitic species are closely related to their host species group, evolving directly from independent ancestors of the host species. However, in the Polyrhachis lama–Diacamma sp. association, the associated species belong to different ant subfamilies. Based on preliminary field surveys, we had presumed that P. lama might have given up its reproductive division of labour, i.e. workers would be able to produce males as well as workers and females parthenogenetically. In this study, this hypothesis was disproved: Polyrhachis lama workers cannot be fertilized and are only able to produce males. In the host–parasite association originating from nests possessing a P. lama queen, workers penetrate surrounding Diacamma sp. nests, carrying brood for rearing within these satellite nests. In this peculiar way, a single P. lama colony is able to exploit several Diacamma sp. colonies simultaneously.     

Nest site selection and induced response in a dominant arboreal ant species

It is well known that arboreal ants, both territorially dominant species and plant ants (e.g., species associated with myrmecophytes or plants housing them in hollow structures), protect their host trees from defoliators. Nevertheless, the presence of an induced defense, suggested by the fact that the workers discovering a leaf wound recruit nestmates, is only known for plant ants. Based on the results from a field study, we show here (1) that colonies of Azteca chartifex, a territorially dominant, neotropical arboreal ant species, mostly selected Goupia glabra (Goupiaceae) trees in which to build their principal carton nests and (2) that plant signals induced workers to recruit nestmates, which patrol the leaves, likely providing the plant with a biotic defense. Furthermore, the number of recruited workers was clearly higher on G. glabra, their most frequently selected host tree species, than on other tree species. These results show that contrary to what was previously believed, induced responses are also found in territorially dominant arboreal ants and so are not limited to the specific associations between myrmecophytes and plant ants.     

Wasps robbing food from ants: a frequent behavior?

Food robbing, or cleptobiosis, has been well documented throughout the animal kingdom. For insects, intrafamilial food robbing is known among ants, but social wasps (Vespidae; Polistinae) taking food from ants has, to the best of our knowledge, never been reported. In this paper, we present two cases involving social wasps robbing food from ants associated with myrmecophytes. (1) Polybioides tabida F. (Ropalidiini) rob pieces of prey from Tetraponera aethiops Smith (Formicidae; Pseudomyrmecinae) specifically associated with Barteria fistulosa Mast. (Passifloraceae). (2) Charterginus spp. (Epiponini) rob food bodies from myrmecophytic Cecropia (Cecropiaceae) exploited by their Azteca mutualists (Formicidae; Dolichoderinae) or by opportunistic ants (that also attack cleptobiotic wasps). We note here that wasps gather food bodies (1) when ants are not yet active; (2) when ants are active, but avoiding any contact with them by flying off when attacked; and (3) through the coordinated efforts of two to five wasps, wherein one of them prevents the ants from leaving their nest, while the other wasps freely gather the food bodies. We suggest that these interactions are more common than previously thought.     

Non-specific association between filamentous bacteria and fungus-growing ants

Fungus-growing ants and their fungal cultivar form a highly evolved mutualism that is negatively affected by the specialized parasitic fungus Escovopsis. Filamentous Pseudonocardia bacteria occurring on the cuticle of attine ants have been proposed to form a mutualistic interaction with these ants in which they are vertically transmitted (i.e. from parent to offspring colonies). Given a strictly vertical transmission of Pseudonocardia, the evolutionary theory predicts a reduced genetic variability of symbionts among ant lineages. The aim of this study was to verify whether actinomycetes, which occur on Acromyrmex octospinosus leaf-cutting ants, meet this expectation by comparing their genotypic variability with restriction fragment length polymorphisms. Multiple actinomycete strains could be isolated from both individual ant workers and colonies (one to seven strains per colony). The colony specificity of actinomycete communities was high: Only 15% of all strains were isolated from more than one colony, and just 5% were present in both populations investigated. Partial sequencing of 16S ribosomal deoxyribonucleic acid of two of the isolated strains assigned both of them to the genus Streptomyces. Actinomycetes could also be isolated from workers of the two non-attine ant species Myrmica rugulosa and Lasius flavus. Sixty-two percent of the strains derived from attine ants and 80% of the strains isolated from non-attine ants inhibited the growth of Escovopsis. Our data suggest that the association between attine ants and their actinomycete symbionts is less specific then previously thought. Soil-dwelling actinomycetes may have been dynamically recruited from the environment (horizontal transmission), probably reflecting an adaptation to a diverse community of microbial pathogens.     

A colony-level response to disease control in a leaf-cutting ant

Parasites and pathogens often impose significant costs on their hosts. This is particularly true for social organisms, where the genetic structure of groups and the accumulation of contaminated waste facilitate disease transmission. In response, hosts have evolved many mechanisms of defence against parasites. Here we present evidence that Atta colombica, a leaf-cutting ant, may combat Escovopsis, a dangerous parasite of Atta's garden fungus, through a colony-level behavioural response. In A. colombica, garden waste is removed from within the colony and transported to the midden - an external waste dump - where it is processed by a group of midden workers. We found that colonies infected with Escovopsis have higher numbers of workers on the midden, where Escovopsis is deposited. Further, midden workers are highly effective in dispersing newly deposited waste away from the dumping site. Thus, the colony-level task allocation strategies of the Atta superorganism may change in response to the threat of disease to a third, essential party.     

Cold resistance depends on acclimation and behavioral caste in a temperate ant

Adjusting to low temperatures is important for animals living in cold environments. We studied the chill–coma recovery time in temperate ant workers (Temnothorax nylanderi) from colonies collected in autumn and spring in Germany. We experimentally acclimated these ant colonies to cold temperatures followed by warm temperatures. As expected, cold-acclimated workers recovered faster from freezing temperatures, but subsequent heat acclimation did not change the short recovery times observed after cold acclimation. Hence, either heat acclimation improves cold tolerance, possibly as a general response to stress, or at least it does not negate enhanced cold tolerance following cold acclimation. Colonies collected in spring showed similar cold tolerance levels to cold-acclimated colonies in the laboratory. Next, we compared the chill–coma recovery time of different worker castes and found that exterior workers recovered faster than interior workers. This difference may be related to their more frequent exposure to cold, higher activity level, or distinct physiology. Interior workers were also heavier and showed a higher gaster-to-head ratio and thorax ratio compared to exterior workers. An obvious difference between exterior and interior workers is activity level, but we found no link between activity and cold tolerance. This suggests that physiology rather than behavioral differences could cause the increased cold tolerance of exterior workers. Our study reveals the importance of acclimation for cold tolerance under natural and standardized conditions and demonstrates differences in cold tolerance and body dimensions in monomorphic behavioral castes of an ant.     

Variable virulence among isolates of Ascosphaera apis: testing the parasite–pathogen hypothesis for the evolution of polyandry in social insects

The queens of many eusocial insect species are polyandrous. The evolution of polyandry from ancestral monoandry is intriguing because polyandry undermines the kin-selected benefits of high intracolonial relatedness that are understood to have been central to the evolution of eusociality. An accumulating body of evidence suggests that polyandry evolved from monoandry in part because genetically diverse colonies better resist infection by pathogens. However, a core assumption of the “parasite–pathogen hypothesis”, that there is variation in virulence among strains of pathogens, remains largely untested in vivo. Here, we demonstrate variation in virulence among isolates of Ascosphaera apis, the causative organism of chalkbrood disease in its honey bee (Apis mellifera) host. More importantly, we show a pathogen–host genotypic interaction for resistance and pathogenicity. Our findings therefore support the parasite-parasite hypothesis as a factor in the evolution of polyandry among eusocial insects.     

The ant genomes have been invaded by several types of mariner transposable elements

To date, only three types of full-length mariner elements have been described in ants, each one in a different genus of the Myrmicinae subfamily: Sinvmar was isolated from various Solenopsis species, Myrmar from Myrmica ruginodis, and Mboumar from Messor bouvieri. In this study, we report the coexistence of three mariner elements (Tnigmar-Si, Tnigmar-Mr, and Tnigmar-Mb) in the genome of a single species, Tapinoma nigerrimum (subfamily Dolichoderinae). Molecular evolutionary analyses of the nucleotide sequence data revealed a general agreement between the evolutionary history of most the elements and the ant species that harbour them, and suggest that they are at the vertical inactivation stage of the so-called Mariner Life Cycle. In contrast, significantly reduced levels of synonymous divergence between Mboumar and Tnigmar-Mb and between Myrmar and Botmar (a mariner element isolated from Bombus terrestris), relative to those observed between their hosts, suggest that these elements arrived to the species that host them by horizontal transfer, long after the species?? split. The horizontal transfer events for the two pairs of elements could be roughly dated within the last 2 million years and about 14 million years, respectively. As would be expected under this scenario, the coding sequences of the youngest elements, Tnigmar-Mb and Mboumar, are intact and, thus, potentially functional. Each mariner element has a different chromosomal distribution pattern according to their stage within the Mariner Life Cycle. Finally, a new defective transposable element (Azteca) has also been found inserted into the Tnigmar-Mr sequences showing that the ant genomes have been invaded by at least four different types of mariner elements.     

The role of tending ants in host plant selection and egg parasitism of two facultative myrmecophilous butterflies

Ovipositing adult females of myrmecophilous lycaenids are expected to select plants based on ant presence in order to maximize the survivorship of immature stages. Usually, larvae feed ants with honey-like solutions and, in turn, ants ward off parasitoids. Nonetheless, a rarely investigated approach is whether ant partners can also extend their protective behavior towards lycaenids eggs. Here, we investigated the ant-related oviposition pattern of Allosmaitia strophius and Rekoa marius; then, we compared egg parasitism according to the presence of ants. Lycaenid oviposition and egg parasitism (in percent) were experimentally compared in ant-present and ant-excluded treatments. The study plant, Heteropterys byrsonimifolia, is an extrafloral nectaried shrub which supports several ant species. We sampled 280 eggs, of which 39.65 % belonged to A. strophius and 60.35 % to R. marius. Both lycaenids eggs were significantly more abundant on branches with ants, especially those with Camponotus crassus and Camponotus blandus, two ant species known to attend to lycaenids. A. strophius and R. marius parasitism was 4.5- and 2.4-fold higher, respectively, in ant-present treatments, but the results were not statistically significant. Our study shows that ant-mediated host plant selection in lycaenids might be much more widespread than previously thought, and not restricted to obligate myrmecophilous species. Tending ants may be inefficient bodyguards of lycaenid eggs, because unlike larvae which release sugared liquids, eggs do not offer obvious rewards to ants. Ants can ward off parasitoids of larvae, as observed elsewhere, but our findings show that positive ant–lycaenid interactions are conditional and depend on immature ontogeny.     

Effect of post-fire resprouting on leaf fluctuating asymmetry, extrafloral nectar quality, and ant–plant–herbivore interactions

Fires in the Cerrado savanna are a severe form of disturbance, but some species are capable of resprouting afterwards. It is unknown, however, how and whether post-fire resprouting represents a stressful condition to plants and how their rapid re-growth influences both the production of biochemical compounds, and interactions with mutualistic ants. In this study, we examined the influence of post-fire resprouting on biotic interactions (ant–plant–herbivore relationships) and on plant stress. The study was performed on two groups of the extrafloral nectaried shrub Banisteriopsis campestris (Malpighiaceae); one group was recovering from fire while the other acted as control. With respect to biotic interactions, we examined whether resprouting influenced extrafloral nectar concentration (milligrams per microliter), the abundance of the ant Camponotus crassus and leaf herbivory rates. Plant stress was assessed via fluctuating asymmetry (FA) analysis, which refers to deviations from perfect symmetry in bilaterally symmetrical traits (e.g., leaves) and indicates whether species are under stress. Results revealed that FA, sugar concentration, and ant abundance were 51.7 %, 35.7 % and 21.7 % higher in resprouting plants. Furthermore, C. crassus was significantly associated with low herbivory rates, but only in resprouting plants. This study showed that post-fire resprouting induced high levels of plant stress and influenced extrafloral nectar quality and ant-herbivore relationships in B. campestris. Therefore, despite being a stressful condition to the plant, post-fire resprouting individuals had concentrated extrafloral nectar and sustained more ants, thus strengthening the outcomes of ant–plant mutualism.     

Resource competition between two fungal parasites in subterranean termites

Subterranean termites live in large groups in underground nests where the pathogenic pressure of the soil environment has led to the evolution of a complex interaction among individual and social immune mechanisms in the colonies. However, groups of termites under stress can show increased susceptibility to opportunistic parasites. In this study, an isolate of Aspergillus nomius Kurtzman, Horn & Hessltine was obtained from a collapsed termite laboratory colony. We determined that it was primarily a saprophyte and, secondarily, a facultative parasite if the termite immunity is undergoing a form of stress. This was determined by stressing individuals of the Formosan subterranean termite Coptotermes formosanus Shiraki via a co-exposure to the virulent fungal parasite Metarhizium anisopliae (Metch.) Sorokin. We also examined the dynamics of a mixed infection of A. nomius and M. anisopliae in a single termite host. The virulent parasite M. anisopliae debilitated the termite immune system, but the facultative, fast growing parasite A. nomius dominated the mixed infection process. The resource utilization strategy of A. nomius during the infection resulted in successful conidia production, while the chance for M. anisopliae to complete its life cycle was reduced. Our results also suggest that the occurrence of opportunistic parasites such as A. nomius in collapsing termite laboratory colonies is the consequence of a previous stress, not the cause of the stress.     

Ant patchiness: a spatially quantitative test in coffee agroecosystems

Arboreal ants form patchy spatial patterns in tropical agroforest canopies. Such patchy distributions more likely occur in disturbed habitats associated with lower ant diversity and resource availability than in forests. Yet, few studies have quantitatively examined these patchy patterns to statistically test if ants are non-randomly distributed or at what scale. Coffee agroecosystems form a gradient of management intensification along which vegetative complexity and ant diversity decline. Using field studies and a spatially explicit randomization model, I investigated ant patchiness in coffee agroecosystems in Chiapas, Mexico varying in management intensity to examine if: (1) coffee intensification affects occurrence of numerically dominant ants, (2) numerical dominants form statistically distinguishable single-species patches in coffee plants, (3) shade trees play a role in patch location, and (4) patch formation or size varies with management intensity. Coffee intensification correlated with lower occurrence frequency of numerically dominant species generally and of one of four taxa examined. All dominant ant species formed patches but only Azteca instabilis was patchy around shade trees. Ant patchiness did vary somewhat with spatial scale and with strata (within the coffee layer vs around shade trees). Patchiness, however, did not vary with management intensity. These results provide quantitative evidence that numerically dominant ants are patchy within the coffee layer at different scales and that shade tree location, but not coffee management intensity, may play a role in the formation of patchy distributions.     

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.     

Knowing your enemies: seasonal dynamics of host–social parasite recognition

Despite its evolutionary significance, behavioural flexibility of social response has rarely been investigated in insects. We studied a host–social parasite system: the slave-making ant Polyergus rufescens and its host Formica rufibarbis. Free-living host workers from parasitized and from unparasitized areas were compared in their level of aggression against the parasite and alien conspecifics. We expected that a seasonal change would occur in the acceptance threshold of F. rufibarbis workers from a parasitized area towards the parasite, whereas F. rufibarbis workers from an unparasitized area would not show substantial changes connected with the parasites peak in activity (raiding and colony-founding season). The results showed a significant adaptive behavioural flexibility of host species workers and are consistent with the acceptance threshold models (Reeve 1989) prediction that recognition systems are not fixed but context-dependent. In particular, host workers from the unparasitized area were highly aggressive towards the parasite regardless of the season, whereas host workers from the parasitized area significantly increased their aggression towards the parasite during its raiding and colony-founding season. Being able to detect and possibly kill a Polyergus scout searching for host nests can be an effective strategy for a Formica colony to avoid being raided or usurped by a parasite queen.     

Out with the garbage: the parasitic strategy of the mantisfly Plega hagenella mass-infesting colonies of the eusocial bee Melipona subnitida in northeastern Brazil

Between April and June of 2012 mantisflies (Plega hagenella) were found to be extensively parasitizing the nests of two groups of managed colonzies of eusocial stingless bees (Melipona subnitida) in the semi-arid region of northeastern Brazil. The mantisfly larvae developed inside closed brood cells of the bee comb, where each mantispid larva fed on the bee larva or pupa present in a single brood cell. Mature mantispid larvae pupated inside silken cocoons spun in place within their hosts' brood cells then emerged as pharate adults inside the bee colony. Pharate adults were never attacked and killed by host colony workers. Instead, colony workers picked up the pharates and removed them from the nest unharmed, treating them similar to the way that the general refuse is removed from the nest. Adult mantispids subsequently eclosed from their pupal exuviae outside the nest. Manipulative experiments showed that post-eclosion adult mantispids placed back within active bee colonies were quickly attacked and killed. These observations demonstrate that pharate and post-eclosion adults of P. hagenella are perceived differently by colony workers and that delayed adult eclosion is an important functional element in the parasitic life strategy of P. hagenella, allowing adults to escape without injury from the bee colonies they parasitize.     

A review of myrmecophily in ant nest beetles (Coleoptera: Carabidae: Paussinae): linking early observations with recent findings

Myrmecophily provides various examples of how social structures can be overcome to exploit vast and well-protected resources. Ant nest beetles (Paussinae) are particularly well suited for ecological and evolutionary considerations in the context of association with ants because life habits within the subfamily range from free-living and predatory in basal taxa to obligatory myrmecophily in derived Paussini. Adult Paussini are accepted in the ant society, although parasitising the colony by preying on ant brood. Host species mainly belong to the ant families Myrmicinae and Formicinae, but at least several paussine genera are not host-specific. Morphological adaptations, such as special glands and associated tufts of hair (trichomes), characterise Paussini as typical myrmecophiles and lead to two different strategical types of body shape: while certain Paussini rely on the protective type with less exposed extremities, other genera access ant colonies using glandular secretions and trichomes (symphile type). We compare these adaptations with other taxonomic groups of insects by joining contemporary research and early sources and discuss the possibility of an attracting or appeasing effect of the secretion. Species that are ignored by their host ants might use chemical mimicry instead. Furthermore, vibrational signals may contribute to ant–beetle communication, and chemical signals have proven to play a role in host finding. The powerful defense chemistry of paussines as “bombardier beetles” is not used in contact with host ants. We attempt to trace the evolution of myrmecophily in paussines by reviewing important aspects of the association between paussine beetles and ants, i.e. morphological and potential chemical adaptations, life cycle, host specificity, alimentation, parasitism and sound production.

Agro-predation: usurpation of attine fungus gardens by Megalomyrmex ants

 A new ant species of Megalomyrmex conducts mass raids to usurp gardens of the fungus-growing ant Cyphomyrmex longiscapus, then lives in the gardens and consumes the cultivated fungus. Unlike attine ants, however, Megalomyrmex sp. does not forage for substrate to manure the gardens; therefore, when gardens become depleted, Megalomyrmex sp. must locate and usurp new gardens. Megalomyrmex sp. workers feed their larvae with attine brood, but only after removing the fungal mycelium that covers the attine larval integument, suggesting that this fungal coat may provide partial protection against other predators. Unlike other known Megalomyrmex species, which coexist as social parasites in attine colonies, Megalomyrmex sp. expels its attine hosts during the garden raids. Megalomyrmex sp. thus maintains a unique agro-predatory lifestyle that is described here for the first time. Received: 18 August 2000 / Accepted in revised form: 1 November 2000