Myrmica ants host highly diverse parasitic communities: from social parasites to microbes

Myrmica ants have been model species for studies in a variety of disciplines, including insect physiology, chemical communication, ant social dynamics, ant population, community ecology, and ant interactions with other organisms. Species belonging to the genus Myrmica can be found in virtually every habitat within the temperate regions of the northern hemisphere and their biology and systematics have been thoroughly studied. These ants serve as hosts to highly diverse parasitic organisms from socially parasitic butterfly caterpillars to microbes, and many Myrmica species even evolved into parasitizing species of their own genus. These parasites have various impacts both on the individuals and on the social structure of their hosts, ranging from morphological malformations to reduction in colony fitness. A comprehensive review of the parasitic organisms supported by Myrmica and the effects of these organisms on individuals and on whole ant colonies has not yet been compiled. Here, we provide a review of the interactions of these organisms with Myrmica ants by discussing host and parasite functional, behavioral or physiological adaptations. In addition, for all “symbiont groups” of Myrmica ants described in this paper, we examine the present limitations of the knowledge at present of their impact on individuals and host colony fitness. In conclusion, we argue that Myrmica ants serve as remarkable resource for the evolution of a wide variety of associated organisms.     

Use of wireless phones and serum β-trace protein in randomly recruited persons aged 18–65 years: a cross-sectional study

Background: There are studies suggesting effects on sleep from pulse-modulated radiofrequency fields used in mobile and cordless phones. So far, reports of adverse effects in observational studies are of limited value for risk assessment while effects from experimental studies seem to be more consistent but unclear as to their importance for health. The aim of this study was to investigate whether use of wireless phones is associated with lower concentrations of β-trace protein (lipocalin-type prostaglandin D synthase), a key enzyme in the synthesis of prostaglandin D₂, an endogenous sleep-promoting neurohormone.

Methods: Three hundred and fourteen people, aged 18–65 years and living in the municipality of Örebro, Sweden, were recruited randomly using the population registry. Total and age-specific linear regression analyses adjusted for known covariates were used to calculate associations between levels of β-trace protein and short- and long-term use of wireless phones.

Results: Overall, no statistically significant association between use of wireless phones and the serum concentration of β-trace protein was found, neither with respect to short-term nor long-term use. Age-specific analyses, however, yielded negative associations for long-term use (cumulative hours of use) and β-trace protein in the youngest age group (18–30 years). Conclusion: This study provided no overall evidence of an association between wireless phone use and serum concentrations of β-trace protein. While the findings in the 18–30 year age group indicating lower concentrations with more cumulative hours of use should be further investigated, no causal inferences can be made from the results of the present study.     

When are ant-attractant devices a worthwhile investment? Vicia faba extrafloral nectaries and Lasius niger ants

Most studies aiming to determine the beneficial effect of ants on plants simply consider the effects of the presence or exclusion of ants on plant yield. This approach is often inadequate, however, as ants interact with both non-tended herbivores and tended Homoptera. Moreover, the interaction with these groups of organisms is dependent on ant density, and these functional relationships are likely to be non-linear. A model is presented here that segregates plant herbivores into two categories depending on the sign of their numerical response to ants (myrmecophiles increase with ants, non-tended herbivores decline). The changes in these two components of herbivores with increasing ant density and the resulting implications for ant-plant mutualisms are considered. It emerges that a wide range of ant densities needs to be considered as the interaction sign (mutualism or parasitism) and strength is likely to change with ant density. The model is used to interpret the results of an experimental study that varied levels of Aphis fabae infestation and Lasius niger ant attendance on Vicia faba bean plants. Increasing ant density consistently reduced plant fitness and thus, in this location, the interaction between the ants and the plant can be considered parasitic. In the Vicia faba system, these costs of ants are unlikely to be offset by other beneficial agents (e.g., parasitoids), which also visit extrafloral nectaries.     

A sophisticated,modular communication contributes to ecological dominance in the invasive ant <Emphasis Type="Italic">Anoplolepis gracilipes</Emphasis>

The ecological success of ants is founded on cooperative behaviour and a well functioning communication. Particularly invasive ants are able to act highly cooperatively, out-compete other species, and become ecologically dominant. Since ant communication is to a large extent chemical, we investigated the pheromone functions involved in foraging and alarm behaviour of the invasive tropical formicine Anoplolepis gracilipes. Our results suggest that long-lasting orientation cues are located in hindguts, while Dufour glands contain short-term attractants that trigger an effective recruitment. Poison gland effects were intermediate between hindgut and Dufour gland in terms of orientation, attraction and longevity. In contrast to the other pheromone sources, mandibular glands have a repellent effect and are most likely involved in alarm behaviour. Taken together, the pheromone glands of A. gracilipes contain functionally distinct signals with considerable differences in persistence. In this respect, its communication is exceptional in formicine ants. A strikingly similar communication system was previously detected in Paratrechina longicornis, another opportunistic and invasive formicine ant. Based on these similarities and the differences compared to non-invasive formicine ants, we discuss the role of chemical signals for the coordination of efficient foraging. We conclude that a sophisticated communication system can contribute significantly to ecological dominance and invasive success, in concert with other well known traits.     

The effect size of aphid‐tending ants in an agricultural tri‐trophic system

Most studies regarding ant–aphid interactions focus only on the direct effects of ants on tended aphids and aphidophagous predators, or the indirect effects on the host plant. Studies evaluating the effects of aphid‐tending ants on more than one trophic level are rare and evaluate only the presence or absence of such effects. Here we assessed the effect sizes of ants in a tri‐trophic system (common bean plants, aphids and lacewing larvae). We tested if the presence of aphid‐tending ants has positive effects on aphid abundance and host‐plant production and negative effects on aphid predator abundance. We also hypothesized that aphid‐tending ants affect more intensely trophic levels that are more directly related to them (i.e., first aphids, then aphid predators and then host plants). We tested these hypotheses in field mesocosms experiments using the presence and absence of ants. We found that aphid‐tending ants have great positive effects on final aphid abundance. Ants also positively affected the number of seeds; however, it was not possible to measure the effect size for this trophic level. Furthermore, ants had negative effects on lacewing larvae only at first release. The effect size of ants was greater for aphids, followed by lacewing larvae, and with no effects on the number of seeds produced. Ants positively affect aphids and host‐plant production, probably by way of honeydew collection preventing the development of entomophagous/saprophytic fungi. On the other hand, ants negatively affect lacewing larvae by excluding them from the host plant. In natural systems, several ant species may attend aphids, differently affecting the organisms of the various trophic levels within the ant–aphid interaction, thereby obscuring the real effect size of ants. Assessing the effect size of aphid‐tending ants on the organisms involved in ant–aphid interactions provides more realistic information about the effects of this interaction on natural systems.     

Trade-offs in group living: transmission and disease resistance in leaf-cutting ants

Sociality can be associated with significant costs due to the increased risk of disease transmission. However, in some organisms the costs may be offset by benefits due to improvements in defences against parasites. To examine this possible trade-off between infection risk and disease resistance, we used Acromyrmex leaf-cutting ants and the entomopathogenic fungus Metarhizium anisopliae as the model system. Ants exposed to the parasite were found to have substantially improved survival when they were kept with nest-mates, while the cost of being in a group in terms of increased disease transmission was very low. The efficiency of transmission is described by the transmission parameter, which decreased with increasing host density showing that transmission rates are inversely density dependent. Both grooming and antibiotic secretions appeared to be important in resistance against the parasite, with the defences of small workers being particularly effective. The results indicate that leaf-cutting ant colonies may have much greater resistance to disease than would be predicted from the high densities of host individuals within them. Unlike most organisms, group living in these ants may actually be associated with a net benefit in terms of disease dynamics.     

Stability and Responsiveness in a Self-Organized Living Architecture

Robustness and adaptability are central to the functioning of biological systems, from gene networks to animal societies. Yet the mechanisms by which living organisms achieve both stability to perturbations and sensitivity to input are poorly understood. Here, we present an integrated study of a living architecture in which army ants interconnect their bodies to span gaps. We demonstrate that these self-assembled bridges are a highly effective means of maintaining traffic flow over unpredictable terrain. The individual-level rules responsible depend only on locally-estimated traffic intensity and the number of neighbours to which ants are attached within the structure. We employ a parameterized computational model to reveal that bridges are tuned to be maximally stable in the face of regular, periodic fluctuations in traffic. However analysis of the model also suggests that interactions among ants give rise to feedback processes that result in bridges being highly responsive to sudden interruptions in traffic. Subsequent field experiments confirm this prediction and thus the dual nature of stability and flexibility in living bridges. Our study demonstrates the importance of robust and adaptive modular architecture to efficient traffic organisation and reveals general principles regarding the regulation of form in biological self-assemblies.     

Bottom-up effects of host-plant species diversity and top-down effects of ants interactively increase plant performance

While plant diversity is well known to increase primary productivity, whether these bottom-up effects are enhanced by reciprocal top-down effects from the third trophic level is unknown. We studied whether pine tree species diversity, aphid-tending ants and their interaction determined plant performance and arthropod community structure. Plant diversity had a positive effect on aphids, but only in the presence of mutualistic ants, leading to a threefold greater number of both groups in the tri-specific cultures than in monocultures. Plant diversity increased ant abundance not only by increasing aphid number, but also by increasing ant recruitment per aphid. The positive effect of diversity on ants in turn cascaded down to increase plant performance; diversity increased plant growth (but not biomass), and this effect was stronger in the presence of ants. Consequently, bottom-up effects of diversity within the same genus and guild of plants, and top-down effects from the third trophic level (predatory ants), interactively increased plant performance.     

Disturbance‐modulated symbioses in termitophily

Symbiosis, the living‐together of unlike organisms, underlies every major transition in evolution and pervades most ecological dynamics. Among examples of symbioses, the simultaneous occupation of a termite nest by its builder termites and intruding invertebrate species (so‐called termitophily) provides suitable macroscopic scenarios for the study of species coexistence in confined environments. Current evidence on termitophily abounds for dynamics occurring at the interindividual level within the termitarium, but is insufficient for broader scales such as the community and the landscape. Here, we inspect the effects of abiotic disturbance on termitophile presence and function in termitaria at these broader scales. To do so, we censused the termitophile communities inhabiting 30 termitaria of distinct volumes which had been exposed to increasing degrees of fire‐induced disturbance in a savanna‐like ecosystem in southeastern Brazil. We provide evidence that such an abiotic disturbance can ease the living‐together of termitophiles and termites. Putative processes facilitating these symbioses, however, varied according to the invader. For nonsocial invaders, disturbance seemed to boost coexistence with termites via the habitat amelioration that termitaria provided under wildfire, as suggested by the positive correlation between disturbance degree and termitophile abundance and richness. As for social invaders (ants), disturbance seemed to enhance associational defenses with termites, as suggested by the negative correlation between the presence of ant colonies and the richness and abundance of other termitarium‐cohabiting termitophiles. It is then apparent that disturbance‐modulated distinct symbioses in these termite nests.     

The Antsy Social Network: Determinants of Nest Structure and Arrangement in Asian Weaver Ants

Asian weaver ants (Oecophylla smaragdina) are arboreal ants that are known to form mutualistic complexes with their host trees. They are eusocial ants that build elaborate nests in the canopy in tropical areas. A colony comprises of multiple nests, usually on multiple trees, and the boundaries of the colony may be difficult to identify. However, they provide the ideal model for studying group living in invertebrates since there are a definite number of nests for a given substrate, the tree. Here, we briefly examine the structure of the nests and the processes involved in the construction and maintenance of these nests. We have described the spatial arrangement of weaver ant nests on trees in two distinct tropical clusters, a few hundred kilometres apart in India. Measurements were made for 13 trees with a total of 71 nests in the two field sites. We have considered a host of biotic and abiotic factors that may be crucial in determining the location of the nesting site by Asian weaver ants. Our results indicate that tree characteristics and architecture followed by leaf features help determine nest location in Asian weaver ants. While environmental factors may not be as influential to nest arrangement, they seem to be important determinants of nest structure. The parameters that may be considered in establishing the nests could be crucial in picking the evolutionary drivers for colonial living in social organisms.     

Who likes it hot? A global analysis of the climatic,ecological, and evolutionary determinants of warming tolerance in ants

Effects of climate warming on wild populations of organisms are expected to be greatest at higher latitudes, paralleling greater anticipated increases in temperature in these regions. Yet, these expectations assume that populations in different regions are equally susceptible to the effects of warming. This is unlikely to be the case. Here, we develop a series of predictive models for physiological thermal tolerances in ants based on current and future climates. We found that tropical ants have lower warming tolerances, a metric of susceptibility to climate warming, than temperate ants despite greater increases in temperature at higher latitudes. Using climatic, ecological and phylogenetic data, we refine our predictions of which ants (across all regions) were most susceptible to climate warming. We found that ants occupying warmer and more mesic forested habitats at lower elevations are the most physiologically susceptible to deleterious effects of climate warming. Phylogenetic history was also a strong indicator of physiological susceptibility. In short, we find that ants that live in the canopies of hot, tropical forest are the most at risk, globally, from climate warming. Unfortunately this is where many, perhaps most, ant and other species on Earth live.     

Radiofrequency electromagnetic radiation exposure inside the metro tube infrastructure in Warszawa

Abstract

Antennas from various wireless communications systems [e.g. mobile phones base transceiver stations (BTS) and handsets used by passengers, public Internet access, staff radiophone transmitters used between engine-drivers and traffic operators] emitting radiofrequency electromagnetic radiation (RF-EMR) are used inside underground metro public transportation. Frequency-selective exposimetric investigations of RF-EMR exposure inside the metro infrastructure in Warsaw (inside metro cars passing between stations and on platforms) were performed. The statistical parameters of exposure to the E-field were analyzed for each frequency range and for a total value (representing the wide-band result of measurements of complex exposure). The recorded exposimetric profiles showed the dominant RF-EMR sources: handsets and BTS of mobile communication systems (GSM 900 and UMTS 2100) and local wireless Internet access (WiFi 2G). Investigations showed that the GSM 900 system is the dominant source of exposure – BTS (incessantly active) on platforms, and handsets – used by passengers present nearby during the tube drive. The recorded E-field varies between sources (for BTS were: medians – 0.22?V/m and 75th percentile – 0.37?V/m; and for handsets: medians – 0.28?V/m and 75th percentile – 0.47?V/m). Maximum levels (peaks) of exposure recorded from mobile handsets exceeded 10?V/m (upper limit of used exposimeters). Broadband measurements of E-field, including the dominant signal emitted by staff radiophones (151?MHz), showed that the level of this exposure of engine-drivers does not exceed 2.5?V/m.     

Grazing and aridity have contrasting effects on the functional and taxonomic diversity of ants

Ants are one of the most abundant and ubiquitous organisms on Earth and play critical roles in multiple ecosystem services such as seed dispersal and nutrient cycling. Despite this, the effects of climatic and land use stressors on particular species or groups of ants are poorly known. We conducted a regional field survey across 108 locations in south-eastern Australia, using correlation network analysis and structural equation modelling to identify how ants respond to environmental stressors. We found contrasting relationships amongst ants, and aridity, and vertebrate grazing intensity and history. Increasing aridity was associated with reduced ant richness, whereas increasing grazing intensity was associated with greater ant richness directly, and indirectly, via reductions in litter depth and perennial grass density. However, these taxonomically diverse groups of ant species still shared contrasting responses to increasing aridity and grazing intensity. We found strong associations between grazing, aridity and the abundance of Seed Harvesters, weak indirect relationships with Generalist Foragers, but no relationships for Predators or Sugar Feeders. Taken together, our work identifies contrasting relationships amongst grazing, aridity and ants (ant ‘winners’ or ‘losers’) across contrasting ecological contexts. Given that increasing aridity is generally associated with lower grazing intensity, our results suggest that locations with more arid sites will have lower ant richness with fewer Seed Harvesters, whereas more mesic sites with high grazing intensity might increase ant richness, and the abundance of specific ant species. Such knowledge is important if we are to maintain critical ant-mediated functions as Earth becomes drier and grazing intensity increases.     

Ant associations in the Neotropical shrub Turnera subulata (Turneraceae): Costs or benefits to the host plant?

The results of ecological interactions depend on the costs and benefits involved in different ecological contexts. Turnera subulata is a shrubby plant with extrafloral nectaries that are associated with ants. Here, we test the hypotheses that the association between T. subulata and ants results in: (i) positive effects on host plant growth and reproduction; (ii) plant herbivory reduction and (iii) inhibition of the host plant visitation by beneficial organisms. Thirty experimental plots were established in northeastern Brazil, either in association with ants or without ants (N = 15 plots/treatment), with four plants each (total 120 plants). Vegetative growth (plant height and number of leaves), reproductive investment (flowers and fruits), herbivory rates and numbers of beneficial visitors were quantified during all phenological stages of the host plant. Data were analysed using generalized linear mixed models. At the host plant maturation stage, we found a trade‐off between growth and reproduction. Plants with ants had lower mean height; however, they invested more in reproduction (a higher number of flowers and fruits) compared to plants without ants. During the flowering stage, the abundance of sucking herbivores was higher in plots without ants but chewing herbivore abundance increased in the maturation stage in plots with ants. The cumulative proportion of leaves with herbivore damage did not differ between treatments, and the presence of ants reduced the number of beneficial visitors (e.g. pollinators and natural enemies) to the host plants. Our results show that association with ants results in some costs for the host plant, however, these costs appear to be offset by the defensive role of ants, which favours plant reproductive investment. In general, our results help to elucidate mechanisms involved in trophic interactions within the complex network of interactions involving ants and plants.     

Are the conformational dynamics and the ligand binding properties of myoglobin affected by exposure to microwave radiation?

The global uptake of mobile communication emphasizes the question about possible adverse consequences of the exposure to low-level radiofrequency radiation from mobile phones on human health as result of so-called "non-thermal effects". In order to state safety guidelines it seems appropriate to start by excluding, if possible, non-specific effects on structural and dynamic properties of fundamental biomolecules such as proteins. Proteins are flexible polyelectrolytes; thus, they are susceptible, in principle, to the action of electromagnetic fields. In this article, we investigated the effects of microwaves on structural and functional properties of Tunnus tynnus myoglobin at 1.95 GHz, a frequency used by new wireless microwave communication systems. The protein solution was exposed for 2.5 h to 51 mW/g SAR (specific absorption rate) level. Measurements of absorption spectroscopy, circular dichroism and fluorescence emission decay in the frequency domain do not exhibit any influence of the radiation on the native structural state of protein macromolecules.     

Thermoregulatory trade-offs result from vegetation removal by a harvester ant

Abstract.  1. Analyses of ecological trade-offs help to explain how organisms balance competing demands. Harvester ants ( Pogonomyrmex occidentalis ), are conspicuous residents of shortgrass prairie in western North America; worker P. occidentalis actively clear all vegetation from the immediate vicinity of their large gravel mounds. This study is based on the prediction that vegetation clearing yields a thermal trade-off by increasing soil temperatures; during cool periods the resulting increase in soil temperature opens new time windows for activity, while during hot periods the soil temperature is more likely to exceed the maximum thermal tolerance for this species. To test the hypothesis that daily and seasonal trade-offs in ant activity result from vegetation removal, the effects of experimentally applied shade on activity patterns were measured.
2. Harvester ant activity correlated highly with ground temperature; experimental shading of ant mounds shifted daily activity patterns by lowering ground temperature. Shading in the morning significantly delayed the onset of ant activity by preventing solar warming of the mound. In contrast, mid-day experimental shading prevented elevation of ground temperatures to above 50 °C and allowed ants to remain active when lethally high temperatures would normally force them inside the mound.
3. A model derived from field data predicted surface ground temperature (and therefore ant activity) based on air temperature and solar radiation, under conditions of sun and shade. For each of six seasons modelled, shade removal yielded a net gain of activity time. These results indicate that vegetation removal by harvester ants produces an advantageous thermoregulatory effect by helping to maximise activity time.     

Recombination,chromosome number and eusociality in the Hymenoptera

Extraordinarily high rates of recombination have been observed in some eusocial species. The most popular explanation is that increased recombination increases genetic variation among workers, which in turn increases colony performance, for example by increasing parasite resistance. However, support for the generality of higher recombination rates among eusocial organisms remains weak, due to low sample size and a lack of phylogenetic independence of observations. Recombination rate, although difficult to measure directly, is correlated with chromosome number. As predicted, several authors have noted that chromosome numbers are higher among the eusocial species of Hymenoptera (ants, bees and wasps). Here, we present a formal comparative analysis of karyotype data from 1567 species of Hymenoptera. Contrary to earlier studies, we find no evidence for an absolute difference between chromosome number in eusocial and solitary species of Hymenoptera. However, we find support for an increased rate of chromosome number change in eusocial taxa. We show that among eusocial taxa colony size is able to explain some of the variation in chromosome number: intermediate‐sized colonies have more chromosomes than those that are either very small or very large. However, we were unable to detect effects of a number of other colony characteristics predicted to affect recombination rate – including colony relatedness and caste number. Taken together, our results support the view that a eusocial lifestyle has led to variable selection pressure for increased recombination rates, but that identifying the factors contributing to this variable selection will require further theoretical and empirical effort.     

Biological responses of mobile phone frequency exposure

Existence of low level electromagnetic fields in the environment has been known since antiquity and their biological implications are noted for several decades. As such dosimetry of such field parameters and their emissions from various sources of mass utilization has been a subject of constant concern. Recent advancement in mobile communications has also drawn attention to their biological effects. Hand held children and adults alike generally use mobile sources as cordless phones in various positions with respect to the body. Further, an increasing number of mobile communication base stations have led to wide ranging concern about possible health effects of radiofrequency emissions. There are two distinct possibilities by which health could be affected as a result of radio frequency field exposure. These are thermal effects caused by holding mobile phones close to the body and extended conversations over a long period of time. Secondly, there could be possibly non thermal effects from both phones and base stations whereby the affects could also be cumulative. Some people may be adversely affected by the environmental impact of mobile phone base stations situated near their homes, schools or any other place. In addition to mobile phones, appliances like microwave oven etc are also in increasing use. Apart from the controversy over the possible health effects due to the non-thermal effect of electromagnetic fields the electromagnetic interaction of portable radio waves with human head needs to be quantitatively evaluated. Relating to this is the criteria of safe exposure to the population at large. While a lot of efforts have gone into resolving the issue, a clear picture has yet to emerge. Recent advances and the problems relating to the safety criteria are discussed.     

Environmental quality mediates the ecological dominance of cooperatively breeding birds

Although social species as diverse as humans and ants are among the most abundant organisms on Earth, animals cooperate and form groups for many reasons. How these different reasons for grouping affect a species' ecological dominance remains unknown. Here we use a theoretical model to demonstrate that the different fitness benefits that animals receive by forming groups depend on the quality of their environment, which in turn impacts their ecological dominance and resilience to global change. We then test the model's key predictions using phylogenetic comparative analysis of >6500 bird species. As predicted, we find that cooperative breeders occurring in harsh and fluctuating environments have larger ranges and greater abundances than non-cooperative breeders, but cooperative breeders occurring in benign and stable environments do not. Using our model, we further show that social species living in harsh and fluctuating environments will be less vulnerable to climate change than non-social species.     

Rethinking edge effects: the unaccounted role of geometric constraints

Edge effects strongly affect the abundance and distribution of organisms across landscapes, with wide‐ranging implications in ecology and conservation biology. The extensive literature on the subject has traditionally considered that edge effects result from the active avoidance or preference of organisms for certain portions of the habitat patch, assuming that abundance is uniform across a patch when environmental conditions are uniform. We demonstrate that this assumption is incorrect due to the so‐far ignored ‘geometric edge effect’ (GEE). In the absence of environmental gradients, abundance of any organism living in a bounded habitat patch will tend to be lower in areas located near the edges compared to areas in the centre of the patch, simply because the areas in the centre receive individuals from all directions, whereas areas near the edge do not receive individuals from outside the patch. This geometric effect was already known for species richness at large geographic scales, the mid‐domain effect, but its importance in the literature of edge effects remained neglected so far. Using simulations, we show that the GEE tends to reduce population abundance and community richness near the edges of bounded habitat patches, and that apparently neutral or negative responses to the edge may occur even when habitat quality is higher near the edges. A published study that detected significant edge effects is reanalyzed, demonstrating that interpreting observed abundance patterns without taking the GEE into account – as traditionally done in the vast literature on edge effects – could provide misleading conclusions. The incorporation of the GEE into sampling and analytical protocols of future studies could advance substantially our ability to understand and predict edge effects in heterogeneous landscapes.