Trophic position and seasonal changes in the diet of the red wood ant Formica aquilonia as indicated by stable isotope analysis

1. Red wood ants are among the most numerous generalist predators and strongly affect the composition of arthropod communities in forest ecosystems. However, their trophic position remains poorly understood. Stable isotope analysis was applied to study the trophic position of Formica aquilonia and reveal seasonal changes in its trophic links with both myrmecophilous aphids and other invertebrates in a mixed forest of western Siberia. 2. The δ¹⁵N values of F. aquilonia exceeded those of herbivores and aphids by approximately 3.5‰. Despite obligate trophobiotic relationships with aphids, F. aquilonia occupied the trophic position of first‐order predator. The higher content of ¹³C in the worker ants, compared with members of grazing food chains, was explained by their consumption of ¹³C‐enriched aphid honeydew. 3. Myrmecophilous tree‐dwelling aphids were enriched in ¹³C and ¹⁵N relative to grass‐inhabiting species, and the honeydew of tree‐dwelling aphids had higher δ¹³C values than those of the honeydew of grass‐inhabiting aphids. 4. The decrease in δ¹³C values of the worker ants from spring and summer to autumn apparently reflected the transition from the collection of tree sap and feeding on the aphid honeydew from trees with high ¹³C content in the spring and early summer to a more diverse liquid diet in late summer, which included ¹³C‐depleted honeydew of aphids from herbs. 5. The prevalence of the ¹⁵N‐depleted aphid honeydew in the ants' diet in the second half of the summer is discussed as one possible explanation for the seasonal decline in δ¹⁵N values of the worker ants.     

Aphid‐tending ants on introduced fennel: can resources derived from non‐native plants alter the trophic position of higher‐order consumers?

1. Although plant invasions often reduce insect abundance and diversity, non‐native plants that support phytophagous insects can subsidise higher trophic levels via elevated herbivore abundance. 2. Here ant–aphid interactions on non‐native fennel on Santa Cruz Island, California are examined. Fennel hosts abundant, honeydew‐producing fennel aphids. The patchiness of fennel and the relative lack of honeydew‐producing insects on other plants at our study sites suggest that assimilation of fennel‐derived honeydew would increase the abundance and decrease the trophic position of the omnivorous, aphid‐tending Argentine ant. 3. To assess the strength of the ant–aphid interaction, a comparison of ant abundance on and adjacent to fennel prior to and 3 weeks after experimental aphid removal was performed. Compared with control plants with aphids, ants declined in abundance on and around fennel plants following aphid removal. At the habitat scale, pitfall traps in fennel‐dominated habitats captured more ants than in fennel‐free scrub habitats. 4. To determine if assimilation of aphid‐produced honeydew reduces the ant's trophic position, variation in δ¹⁵N values among ants, plants and other arthropods was analysed. Unexpectedly, δ¹⁵N values for ants in fennel‐dominated habitats were higher than those of arthropod predators from the same sites and also higher than those of ants from fennel‐free habitats. 5. Our results illustrate how introduced plants that support phytophagous insects appear to transfer energy to higher trophic levels via elevated herbivore abundance. Although assimilation of fennel‐derived honeydew did not appear to reduce consumer trophic position, spatial variation in alternative food resources might obscure contributions from honeydew.     

Plant‐derived differences in the composition of aphid honeydew and their effects on colonies of aphid‐tending ants

In plant–ant–hemipteran interactions, ants visit plants to consume the honeydew produced by phloem‐feeding hemipterans. If genetically based differences in plant phloem chemistry change the chemical composition of hemipteran honeydew, then the plant's genetic constitution could have indirect effects on ants via the hemipterans. If such effects change ant behavior, they could feed back to affect the plant itself. We compared the chemical composition of honeydews produced by Aphis nerii aphid clones on two milkweed congeners, Asclepias curassavica and Asclepias incarnata, and we measured the responses of experimental Linepithema humile ant colonies to these honeydews. The compositions of secondary metabolites, sugars, and amino acids differed significantly in the honeydews from the two plant species. Ant colonies feeding on honeydew derived from A. incarnata recruited in higher numbers to artificial diet, maintained higher queen and worker dry weight, and sustained marginally more workers than ants feeding on honeydew derived from A. curassavica. Ants feeding on honeydew from A. incarnata were also more exploratory in behavioral assays than ants feeding from A. curassavica. Despite performing better when feeding on the A. incarnata honeydew, ant workers marginally preferred honeydew from A. curassavica to honeydew from A. incarnata when given a choice. Our results demonstrate that plant congeners can exert strong indirect effects on ant colonies by means of plant‐species‐specific differences in aphid honeydew chemistry. Moreover, these effects changed ant behavior and thus could feed back to affect plant performance in the field.     

Elevated temperatures alter an ant‐aphid mutualism

Ant‐hemipteran mutualisms are keystone interactions that can be variously affected by warming: these mutualisms can be strengthened or weakened, or the species can transition to new mutualist partners. We examined the effects of elevated temperatures on an ant‐aphid mutualism in the subalpine zone of the Rocky Mountains in Colorado, USA. In this system, inflorescences of the host plant, Ligusticum porteri Coult. & Rose (Apiaceae), are colonized by the ant‐tended aphid Aphis asclepiadis Fitch or less frequently by the non‐ant tended aphid Cavariella aegopodii (Scopoli) (both Hemiptera: Aphididae). Using an 8‐year observational study, we tested for two key mechanisms by which ant‐hemipteran mutualisms may be altered by climate change: shifts in species identity and phenological mismatch. Whereas the aphid species colonizing the host plant is not changing in response to year‐to‐year variation in temperature, we found evidence that a phenological mismatch between ants and aphids could occur. In warmer years, colonization of host plant inflorescences by ants is decreased, whereas for A. asclepiadis aphids, host plant colonization is mostly responsive to date of snowmelt. We also experimentally established A. asclepiadis colonies on replicate host plants at ambient and elevated temperatures. Ant abundance did not differ between aphid colonies at ambient vs. elevated temperatures, but ants were less likely to engage in tending behaviors on aphid colonies at elevated temperatures. Sugar composition of aphid honeydew was also altered by experimental warming. Despite reduced tending by ants, aphid colonies at elevated temperatures had fewer intraguild predators. Altogether, our results suggest that higher temperatures may disrupt this ant‐aphid mutualism through both phenological mismatch and by altering benefits exchanged in the interaction.     

Costs and benefits of ant attendance to the drepanosiphid aphid Tuberculatus quercicola

The defensive effects of ants against aphid predators have been well documented in the mutualistic relationship of aphids and their attending ants. However, it is not clear whether ant attendance has any direct effect on the aphids' growth and reproduction. Through field experiments, this study evaluates the benefits and, in particular, the costs of ant attendance to aphid colonies, focusing on the drepanosiphid aphid Tuberculatus quercicola which is associated with the Daimyo oak, Quercus dentata , and which is always attended by the red wood ant Formica yessensis . Ant attendance was clearly beneficial to the aphid; the exclusion of ants led to a significant increase in the extinction rate of aphid colonies. However, MANOVA and randomized block ANOVA indicated that in colonies continuously attended by ants, aphids had significantly smaller body size and produced a smaller number of embryos than in colonies isolated from ants when they were reared under homogeneous host conditions free from natural enemies. Thus, ant attendance had a negative influence on the growth and reproduction of the aphids, even though it contributed to the greater longevity of the aphid colonies. We hypothesize that ant-attended aphids are under intense selective pressures that act against aphid clones which fail to attract many ants, so that aphids have developed an adaptive mechanism to allocate a larger fraction of resources to the honeydew when they are requested to do so by the ants in order to ensure the ants' consistent visitation.     

Influence of macronutrient imbalance on native ant foraging and interspecific interactions in the field

1. Ants interact with a diversity of organisms. These interactions, coupled with their abundance, cause ants to have ecologically important effects across multiple trophic levels. 2. Empirical study of ant nutritional ecology has led to the prediction that a macronutrient imbalance will affect ant behaviour and interspecific interactions that underlie these broad‐scale effects. Excess carbohydrate relative to protein is predicted to increase ant aggressiveness, predatory tendency and foraging activity, and to decrease collection of hemipteran honeydew and plant nectar. 3. In field experiments conducted in 2009 and 2010, captive colony fragments of a native ant, Formica podzolica (Hymenoptera: Formicidae), were provided with either simulated prey or carbohydrate solution ad libitum. Foraging behaviours and interactions with flowers, myrmecophilous aphids and aphid natural enemies on wild‐grown plants were documented. 4. Strong effects of macronutrient imbalance on foraging manifested quickly and consistently across colonies; in accordance with predictions, prey‐fed foragers collected both honeydew and floral nectar, whereas carbohydrate‐fed ants ceased collecting these resources. Counter to predictions, carbohydrate‐fed ants dramatically lowered their activity levels and did not prey upon aphids. 5. Ants had no effect on aphid enemies in 2009, when the latter were relatively rare, but decreased their abundance in 2010. Despite this protection, the net effect of ants on aphids was negative (measured only in 2009). Prey‐fed ants demonstrated a strong preference for honeydew over floral nectar, thus demonstrating that a macronutrient imbalance may lead to different interactions with similar resources. 6. This study links ant nutrition and community ecology by demonstrating the rapid, asymmetric and multitrophic consequences of nutritionally mediated behaviour.     

Introduced ants reduce interaction diversity in a multi‐species,ant–aphid mutualism

Mutualisms contribute in fundamental ways to the origin, maintenance and organization of biological diversity. Introduced species commonly participate in mutualisms, but how this phenomenon affects patterns of interactions among native mutualists remains incompletely understood. Here we examine how networks of interactions among aphid‐tending ants, ant‐tended aphids, and aphid‐attacking parasitoid wasps differ between 12 spatially paired riparian study sites with and without the introduced Argentine ant Linepithema humile in southern California. To resolve challenges in species identification, we used DNA barcoding to identify aphids and screen for parasitoid wasps (developing inside their aphid hosts) from 170 aphid aggregations sampled on arroyo willow Salix lasiolepis. Compared to uninvaded sites, invaded sites supported significantly fewer species of aphid‐tending ants and ant‐tended aphids. At invaded sites, for example, we found only two species of ant‐tended aphids, which were exclusively tended by L. humile, whereas at uninvaded sites we found 20 unique ant–aphid interactions involving eight species of ant‐tended aphids and nine species of aphid‐tending ants. Ant–aphid linkage density was thus significantly lower at invaded sites compared to uninvaded sites. We detected aphid parasitoids in 14% (28/198) of all aphid aggregations. Although the level of parasitism did not differ between invaded and uninvaded sites, more species of wasps were detected within uninvaded sites compared to invaded sites. These results provide a striking example of how the assimilation of introduced species into multi‐species mutualisms can reduce interaction diversity with potential consequences for species persistence.     

Trophic ecology of the invasive argentine ant: spatio-temporal variation in resource assimilation and isotopic enrichment

Studies of food webs often employ stable isotopic approaches to infer trophic position and interaction strength without consideration of spatio-temporal variation in resource assimilation by constituent species. Using results from laboratory diet manipulations and monthly sampling of field populations, we illustrate how nitrogen isotopes may be used to quantify spatio-temporal variation in resource assimilation in ants. First, we determined nitrogen enrichment using a controlled laboratory experiment with the invasive Argentine ant (Linepithema humile). After 12 weeks, worker δ¹⁵N values from colonies fed an animal-based diet had δ¹⁵N values that were 5.51% greater compared to colonies fed a plant-based diet. The shift in δ¹⁵N values in response to the experimental diet occurred within 10 weeks. We next reared Argentine ant colonies with or without access to honeydew-producing aphids and found that after 8 weeks workers from colonies without access to aphids had δ¹⁵N values that were 6.31% larger compared to colonies with access to honeydew. Second, we sampled field populations over a 1-year period to quantify spatio-temporal variability in isotopic ratios of L. humile and those of a common native ant (Solenopsis xyloni). Samples from free-living colonies revealed that fluctuations in δ¹⁵N were 1.6–2.4‰ for L. humile and 1.8–2.9‰ for S. xyloni. Variation was also detected among L. humile castes: time averaged means of δ¹⁵N varied from 1.2 to 2.5‰ depending on the site, with δ¹⁵N values for queens ≥ workers > brood. The estimated trophic positions of L. humile and S. xyloni were similar within a site; however, trophic position for each species differed significantly at larger spatial scales. While stable isotopes are clearly useful for examining the trophic ecology of arthropod communities, our results suggest that caution is warranted when making ecological interpretations when stable isotope collections come from single time periods or life stages.     

Do aphids actively search for ant partners?

The aphid–ant mutualistic relationships are not necessarily obligate for neither partners but evidence is that such interactions provide them strong advantages in terms of global fitness. While it is largely assumed that ants actively search for their mutualistic partners namely using volatile cues; whether winged aphids (i.e., aphids’ most mobile form) are able to select ant‐frequented areas had not been investigated so far. Ant‐frequented sites would indeed offer several advantages for these aphids including a lower predation pressure through ant presence and enhanced chances of establishing mutuaslistic interactions with neighbor ant colonies. In the field, aphid colonies are often observed in higher densities around ant nests, which is probably linked to a better survival ensured by ants’ services. Nevertheless, this could also result from a preferential establishment of winged aphids in ant‐frequented areas. We tested this last hypothesis through different ethological assays and show that the facultative myrmecophilous black bean aphid, Aphis fabae L., does not orientate its search for a host plant preferentially toward ant‐frequented plants. However, our results suggest that ants reduce the number of winged aphids leaving the newly colonized plant. Thus, ants involved in facultative myrmecophilous interactions with aphids appear to contribute to structure aphid populations in the field by ensuring a better establishment and survival of newly established colonies rather than by inducing a deliberate plant selection by aphid partners based on the proximity of ant colonies.     

How an ant decides to prey on or to attend aphids

The following results on the behavior decision making of the antLasius niger toward two species of myrmecophilous aphidsLachnus tropicalis andMyzocallis kuricola on chestnut trees have been found. (1) An individual worker consistently attended only one aphid species, even if her nestmates attended other aphid species on the same tree. (2) The ants preyed less on the aphid species which they attended than on other myrmecophilous aphid species. (3) The ants preyed less on the aphids which had been attended by their nestmates, even if both aphids were the same species. (4) The ants preyed less on aphids which had provided honeydew to their nestmates. (5) The increased aphid density per ant led to an increase in the rate of predation on the introduced aphids by the ants. These results suggest that each worker ofL. niger chooses aphid species to attend from her experience. In addition, the workers can recognize whether an aphid has been attended by their nestmates and whether an aphid has given their nestmates honeydew. Through these processes, each worker decides to attend or to prey on the aphid. As a result, they may realize efficient collective foraging dependent on aphid density per worker.     

Alate production in an aphid in relation to ant tending and alarm pheromone

1. Winged dispersal is vital for aphids as predation pressure and host plant conditions fluctuate. 2. Ant‐tended aphids also need to disperse, but this may represent a cost for the ants, resulting in an evolutionary conflict of interest over aphid dispersal. 3. The combined effects of aphid alarm pheromone, indicating predation risk, and ant attendance on the production of winged aphids were examined in an experiment with Aphis fabae (Homoptera: Aphididae) (Scopoli 1763) aphids and Lasius niger (Formicidae: Formicinae) (Linné, 1758) ants. 4. This study is the first to investigate the joint effects of alarm pheromone and ant attendance, and also the first to detect an influence of alarm pheromone on the production of winged morphs in A. fabae. 5. After a period of 2 weeks, it was found that aphid colonies exposed to intermittent doses of alarm pheromone produced more winged individuals, whereas ant tending had the opposite effect. The effects were additive on a log scale, and ant attendance had a greater proportional influence than exposure to alarm pheromone. A tentative conclusion is that ants have gained the upper hand in an evolutionary conflict about aphid dispersal.     

Tri-trophic level interactions affect host plant development and abundance of insect herbivores

Understanding the interactions among plants, hemipterans, and ants has provided numerous insights into a range of ecological and evolutionary processes. In these systems, however, studies concerning the isolated direct and indirect effects of aphid colonies on host plant and other herbivores remain rare at best. The aphid Uroleucon erigeronensis forms dense colonies on the apical shoots of the host plant Baccharis dracunculilfolia (Asteraceae). The honeydew produced by these aphids attracts several species of ants that might interfere with other herbivores. Four hypotheses were tested in this system: (1) ants tending aphids reduce the abundance of other herbivores; (2) the effects of ants and aphids upon herbivores differ between chewing and fluid-sucking herbivores; (3) aphids alone reduce the abundance of other herbivores; and (4), the aphid presence negatively affects B. dracunculifolia shoot growth. The hypotheses were evaluated with ant and aphid exclusion experiments, on isolated plant shoots, along six consecutive months. We adjusted linear mixed-effects models for longitudinal data (repeated measures), with nested spatial random effect. The results showed that: (1) herbivore abundance was lower on shoots with aphids than on shoots without aphids, and even lower on shoots with aphids and ants; (2) both chewing and fluid-sucking insects responded similarly to the treatment, and (3) aphid presence affected negatively B. dracunculifolia shoot growth. Thus, since aphids alone changed plant growth and the abundance of insect herbivores, we suggest that the ant–aphid association is important to the organization of the system B. dracunculifolia-herbivorous insects.     

Towards a nutritional ecology of invasive establishment: aphid mutualists provide better fuel for incipient Argentine ant colonies than insect prey

Many potential species invasions fail before establishment. This is likely especially true for invasive Argentine ants that must overcome a severe founding bottleneck and transition from propagules that rely on protein-rich prey to massive supercolonies that dominate by consuming carbohydrate-rich honeydew from hemipteran mutualists. While this dietary shift supports the classic idea that protein fuels early colony development and carbohydrates maintain adult workers, recent evidence suggests that carbohydrates can govern initial colony establishment. In this study, we use lab experiments to show that resources from aphid mutualists had greater benefits for Argentine ant propagule survival, maintenance, growth, and worker activity rates than did prey items. These effects persisted at low aphid densities, and when colonies were otherwise starved. Moreover, prey-starved colonies did not appear to consume aphids, suggesting that carbohydrate-rich honeydew is a mechanism that facilitates colony establishment. Combined, these results support a hypothesis that the dietary shift from prey to honeydew is driven more by increased access to hemipterans after establishment, than by specific benefits of prey early in colony development. The results highlight the important role of nutritional ecology for studying invasive establishment, linking propagule success not only to the supply of food resources, but also to their quality.     

Genetically based population variation in aphid association with ants and predators

A species’ genotype can have extended consequences for the structure of the surrounding community, but few studies have investigated the extended consequences of genetic variation in animals. Accordingly, I examined the importance of genetically based variation among five populations of the ant-tended aphid Aphis asclepiadis for its interactions with both ants and predators. In a common environment, aphid source population accounted for 23 and 17% of the variation in the occurrence of ants and predators, respectively. Ant exclusion increased predator abundance, accounting for 25% of variation, but there was no detectable influence of ants on aphid abundance. There was an indication that aphid source populations varied in honeydew quality, but this was uncorrelated with rates of ant attendance. This study provides the first evidence for genetic variation in aphids for attractiveness to ants, and underscores the important link between intra-specific genetic variation in aphids and the processes governing arthropod community structure.     

The effects of the invasive Argentine ant (Linepithema humile) and the native ant Prenolepis imparis on the structure of insect herbivore communities on willow trees (Salix lasiolepis)

Abstract 1. We examined the relative effects of the invasive Argentine ant, Linepithema humile, and a common native ant, Prenolepis imparis, on the community of herbivorous insects occurring on willow trees, Salix lasiolepis in Northern California, U.S.A. 2. Using paired control and treatment branches from which we excluded ants and other non‐volant predators, we found that effects of Argentine ants on the herbivore community were generally similar to those of P. imparis. Argentine ants and P. imparis suppressed the damage by skeletonising insects by 50%, but had little effect on most other external‐feeding or internal‐feeding guilds. 3. The abundance of aphids was 100% greater in the presence of Argentine ants, but there was no effect on aphid numbers in the presence of P. imparis. Late season aphid numbers were substantially higher in the presence of Argentine ants, but not P. imparis. 4. The effects of Argentine ants on skeletonising insects and aphids combined with the overwhelming abundance of Argentine ant workers, suggests that they may have substantial, but often overlooked, effects on the herbivore communities on other plant species in or near riparian habitats in which they invade.     

Flexibility in the composition and concentration of amino acids in honeydew of the drepanosiphid aphid Tuberculatus quercicola

Abstract 1. Mutualistic interactions between aphids and ants are mediated by honeydew that aphids produce. Previous work showed that when attended by the ant Formica yessensis Forel (Hymenoptera: Formicidae), nymphs of the aphid Tuberculatus quercicola (Matsumura) (Homoptera: Aphididae) developed into significantly smaller adults with lower fecundity than did nymphs that were not ant attended.
2. This study tested the hypothesis that this cost of ant attendance arises through changes in the quality and quantity of honeydew. Ant-attended and ant-excluded aphid colonies were prepared in the field. The composition and concentration of amino acids were compared between the honeydew produced by ant-attended colonies and that produced by ant-excluded colonies.
3. The aphids excreted smaller droplets of honeydew, but also excreted them more frequently, in ant-attended colonies than in ant-excluded colonies. The honeydew of ant-attended aphids contained more types of amino acid, and a significantly higher total concentration of amino acids, than did the honeydew of ant-excluded aphids.
4. These results suggest that the increase in the concentration of amino acids in honeydew leads to a shortage of nitrogen available for aphid growth and reproduction, resulting in lower performance under ant attendance.
5. With the advance of seasons, a significant reduction was found in both the total free amino acid concentration in phloem sap and the frequency of honeydew excretion; however the total concentration of amino acids in the honeydew did not vary significantly during the seasons, suggesting that aphids keep the quality of honeydew constant in order to maintain ant visitation.     

Do exaptations facilitate mutualistic associations between invasive and native species?

As invasive species are key threats to ecosystem structure and function, it is essential to understand the factors underlying their success. Enigmatically, mutualistic organisms are often successful in colonizing novel environments even though they commonly persist only through intricate relationships with other species. Mutualistic ants, for example, protect aphids from natural enemies while collecting carbohydrate–rich honeydew. To facilitate this interaction, ants have evolved aggressive responses to aphid alarm pheromone emissions. As invasive and native mutualists have not evolved together, however, it is unclear if this form of cross-species communication exists between these two parties thereby facilitating these novel interactions. We address this hypothesis by assessing whether the invasive Argentine ant, Linepithema humile, responds to native poplar aphid, Chaitophorus populicola, alarm signals. Here, we show that interspecific signalling does exist in this newly established mutualistic interaction. Argentine ant workers exhibit increased aggression and double the number of visits to an aphid colony after an aphid alarm signal is emitted. We suggest that pre-adaptations may facilitate the emergence of mutualistic associations between many invasive and native species.     

Bacteria may contribute to distant species recognition in ant–aphid mutualistic relationships

Mutualistic interactions between ant and aphid species have been the subject of considerable historical and contemporary investigations, the primary benefits being cleaning and protection for the aphids and carbohydrate‐rich honeydew for the ants. Questions remained, however, as to the volatile semiochemical factor influencing this relationship. A recent study highlighted the role of bacterial honeydew volatile compounds in ant attraction. Here, ant's ability to distantly discriminate 2 aphid species was investigated based on bacterial honeydew semiochemicals emissions using a two‐way olfactometer. Both the mutualistic aphid Aphis fabae L. and the nonmyrmecophilous aphid Acyrthosiphon pisum Harris were found to be attractive for the ant Lasius niger L. The level of attraction was similar in both assays (control vs. one of the aphid species). However, when given a choice between these 2 aphid species, ants showed a significant preference for Aphis fabae. Honeydew volatiles, mostly from bacterial origins, are known to be a key element in ant attraction. Using the same olfactometry protocol, the relative attractiveness of volatiles emitted by honeydews collected from each aphid species and by bacteria isolated from each honeydew was investigated. Again, ants significantly preferred volatiles released by Aphis fabae honeydew and bacteria. This information suggests that microbial honeydew volatiles enable ants to distantly discriminate aphid species. These results strengthen the interest of studying the occurrence and potential impact of microorganisms in insect symbioses.     

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.     

Getting carbohydrates in ants: damage of the young cereal sprouts by ants of the genus Myrmica Latreille, 1804 (Hymenoptera: Formicidae), to obtain plant sap

Summary

Carbohydrate food is of high importance for survival of ant colonies. Ants are known to use sugary excretions (honeydew) of various insects, nectar of floral and extrafloral nectaries, and even sap of some trees. However, the ability of ants to use sap of herbaceous plants has not been mentioned. This is the first evidence that ants of the genus Myrmica can intentionally ‘cut off’ young cereal sprouts to obtain plant sap. The investigation was carried out in a laboratory in 2018 and 2019 and involved three ant species of the genus Myrmica [12 colonies of M. rubra (Linnaeus, 1758); eight colonies of M. ruginodis Nylander, 1846; and five colonies of M. scabrinodis Nylander, 1846]. First observations were made occasionally in 10 ant colonies during the study of ant–aphid interactions. After three days of carbohydrate starvation, ants were supplied with the plants of wheat infested with aphids of Schizaphis graminum (Rondani, 1852). Within the first day in addition to ordinary trophobiotic relations with aphids, the workers of all the studied colonies demonstrated unexpected behaviour: they ‘cut off’ some sprouts and collected sap of these plants. The experimental investigation in 15 ant colonies of various sizes (about 150, 300 and 500 workers) supplied with the plants infested or non-infested with aphids has shown that getting sap of herbs depends greatly on ant colony needs and available resources. The number of damaged plants was much higher both in the larger colonies of ants and in the absence of aphids. This way of getting carbohydrates allows ants to quickly obtain some extra food needed to maintain colony viability and seems to be one of the mechanisms promoting survival of ants in conditions of acute carbohydrate deficiency. At the same time, ants avoid using plant sap when there are more available alternative carbohydrate resources.