Predator and parasitoid attacking ant-attended aphids: effects of predator presence and attending ant species on emerging parasitoid numbers

Interaction between a predator and a parasitoid attacking ant-attended aphids was examined in a system on photinia plants, consisting of the aphid Aphis spiraecola, the two ants Lasius japonicus and Pristomyrmex pungens, the predatory ladybird beetle Scymnus posticalis, and the parasitoid wasp Lysiphlebus japonicus. The ladybird larvae are densely covered with waxy secretion and are never attacked by attending ants. The parasitoid females are often attacked by ants, but successfully oviposit by avoiding ants. The two ants differ in aggressiveness towards aphid enemies. Impacts of the predator larvae and attending ant species on the number of parasitoid adults emerging from mummies per aphid colony were assessed by manipulating the presence of the predator in introduced aphid colonies attended by either ant. The experiment showed a significant negative impact of the predator on emerging parasitoid numbers. This is due to consumption of healthy aphids by the predator and its predation on parasitized aphids containing the parasitoid larvae (intraguild predation). Additionally, attending ant species significantly affected emerging parasitoid numbers, with more parasitoids in P. pungens-attended colonies. This results from the lower extent of interference with parasitoid oviposition by the less aggressive P. pungens. Furthermore, the predator reduced emerging parasitoid numbers more when P. pungens attended aphids. This may be ascribed to larger numbers of the predator and the resulting higher levels of predation on unparasitized and parasitized aphids in P. pungens-attended colonies. In conclusion, a negative effect of the predator on the parasitoid occurs in ant-attended aphid colonies, and the intensity of the interaction is affected by ant species.     

Intraguild interactions between spiders and ants and top-down control in a grassland food web

In most terrestrial ecosystems ants (Formicidae) as eusocial insects and spiders (Araneida) as solitary trappers and hunters are key predators. To study the role of predation by these generalist predators in a dry grassland, we manipulated densities of ants and spiders (natural and low density) in a two-factorial field experiment using fenced plots. The experiment revealed strong intraguild interactions between ants and spiders. Higher densities of ants negatively affected the abundance and biomass of web-building spiders. The density of Linyphiidae was threefold higher in plots without ant colonies. The abundance of Formica cunicularia workers was significantly higher in spider-removal plots. Also, population size of springtails (Collembola) was negatively affected by the presence of wandering spiders. Ants reduced the density of Lepidoptera larvae. In contrast, the abundance of coccids (Ortheziidae) was positively correlated with densities of ants. To gain a better understanding of the position of spiders, ants and other dominant invertebrate groups in the studied food web and important trophic links, we used a stable isotope analysis (¹⁵N and ¹³C). Adult wandering spiders were more enriched in ¹⁵N relative to ¹⁴N than juveniles, indicating a shift to predatory prey groups. Juvenile wandering and web-building spiders showed δ¹⁵N ratios just one trophic level above those of Collembola, and they had similar δ¹³C values, indicating that Collembola are an important prey group for ground living spiders. The effects of spiders demonstrated in the field experiment support this result. We conclude that the food resource of spiders in our study system is largely based on the detrital food web and that their effects on herbivores are weak. The effects of ants are not clear-cut and include predation as well as mutualism with herbivores. Within this diverse predator guild, intraguild interactions are important structuring forces.     

Emergent Impacts of Ant and Spider Interactions: Herbivory Reduction in a Tropical Savanna Tree

Multiple predators often have effects on their common prey populations that cannot be predicted by summing the effects of each predator at a time. When predators forage on the same vegetation substrate, intraguild interactions might cause emergent outcomes for the plants on which the predators co‐occur. We experimentally evaluated the effects of spiders and ants on herbivory and reproduction in the extrafloral nectary‐bearing tree Qualea multiflora (Vochysiaceae). Plants were divided in four experimental groups, depending on the presence or absence of ants and spiders. We compared the effects of each treatment on richness and abundance of chewing and sucking herbivores and on herbivory (leaf area loss). We also evaluated the impact of predators on the production of buds, fruits and seeds, and weight of the fruits. The presence of ants reduced the abundance and richness of spiders, but spiders did not affect the abundance and richness of ants. Only the removal of ants resulted in a significant increase in the abundance of herbivores and herbivore richness. Herbivory, however, was also affected by spiders. In addition, we found a significant interaction effect of ants and spiders on herbivory, indicating an emergent multiple predator effect. Neither ants nor spiders had an impact on the number of buds produced, number of fruits per bud, and seeds per fruits or fruit weight. This study highlights the importance of evaluating the effect of the predator fauna as a whole and not only one specific group on herbivory.     

Effects of the concurrent exclusion of ants and earwigs on aphid abundance in an organic citrus grove

Based on the well-known mutualism between ants (Hymenoptera: Formicidae) and aphids (Homoptera: Aphididae), we conducted a five-year experiment of ant-exclusion from the canopies of citrus trees as a possible method of biological control of aphids. However, our results showed that the exclusion of ants from the canopies increased, instead of reducing, aphid abundance. To explain this unexpected result, we reasoned that the exclusion of ants from the canopies might also have excluded crawling insects that prey on aphids, such as the European earwig (Forficula auricularia L., Dermaptera: Forficulidae). Such a possibility is supported by the negative relationship between aphid density and the abundance of earwigs, consistent with a top-down control of aphids by earwigs. In contrast, the abundance of other aphid predators (Coleoptera: Coccinellidae, and Heteroptera) had no such negative effect on aphid density but a positive one, suggesting a bottom-up control, and showed no differences between control and ant-excluded trees. Thus, the most likely explanation for the increase in aphid abundance in the ant-excluded trees is the absence of earwigs from the canopies of the experimental trees, providing further evidence of the major role that earwigs play as control agents of aphids in cultivated trees.     

The response of an aphid tending ant to artificial extra-floral nectaries on different host plants

Sap-feeding homopterans, which reduce the fitness of their host plants, are often tended by ants that feed on their honeydew. The composition of the honeydew varies with both the aphid and the host plant. Extra-floral nectaries (EFNs) are believed to have evolved to attract attending ants, protecting the hosts, but it is unknown if EFNs on different plants have the same impact on the relations between an aphid species feeding on those plants and its tending ant. Experimental research was conducted to examine the attraction of Tapinoma erraticum scout ants to honeydew from the aphid Aphis gossypii feeding on two different plants, Prunus amygdalus and Mentha piperita, negligence of tending the aphids, and survival of the aphids in the presence of artificial EFNs. The scout ants were significantly more attracted to artificial nectar dispensed on P. amygdalus leaves than on M. piperita, or aphids on both plants and water. They neglected aphids in the presence of artificial EFNs on M. piperita but not on P. amygdalus. The aphid population on M. piperita did not statistically change in the presence of artificial EFNs during the 8 days of the third experiment. On P. amygdalus, the aphids succeeded in developing fully to winged form. In conclusion, the responses of the ants tending aphids to the presence of artificial EFNs were influenced by the host plant.     

Jumping spiders (Salticidae) enhance the seed production of a plant with extrafloral nectaries

Many plants secrete nectar from extrafloral nectaries (EFNs), specialized structures that usually attract ants which can act as plant defenders. We examined the nectar-mediated interactions between Chamaecrista nictitans (Caesalpineaceae) and jumping spiders (Araneae, Salticidae) for 2 years in old fields in New Jersey, USA. Previous research suggests that spiders are entirely carnivorous, yet jumping spiders (Eris sp. and Metaphidippus sp.) on C. nictitans collected nectar in addition to feeding on herbivores, ants, bees, and other spiders. In a controlled-environment experiment, when given a choice between C. nictitans with or without active EFNs, foraging spiders spent 86% of their time on plants with nectar. C. nictitans with resident jumping spiders did set significantly more seed than plants with no spiders, indicating a beneficial effect from these predators. However, the presence of jumping spiders did not decrease numbers of Sennius cruentatus (Bruchidae), a specialist seed predator of C. nictitans. Jumping spiders may provide additional, unexpected defense to plants possessing EFNs. Plants with EFNs may therefore have beneficial interactions with other arthropod predators in addition to nectar-collecting ants. Received: 27 May 1998 / Accepted: 23 December 1998     

Associational resistance or susceptibility: the indirect interaction between chemically‐defended and non‐defended herbivore prey via a shared predator

Many organisms possess chemical defences against their natural enemies, which render them unpalatable or toxic when attacked or consumed. These chemically‐defended organisms commonly occur in communities with non‐ or less‐defended prey, leading to indirect interactions between prey species, mediated by natural enemies. Although the importance of enemy‐mediated indirect interactions have been well documented (e.g. apparent competition), how the presence of prey chemical defences may affect predation of non‐defended prey in terrestrial communities remains unclear. Here, an experimental approach was used to study the predator‐mediated indirect interaction between a chemically‐defended and non‐defended pest aphid species. Using laboratory‐based mesocosms, aphid community composition was manipulated to include chemically‐defended (CD) aphids Brevicoryne brassicae, non‐defended (ND) aphids Myzus persicae or a mixed assemblage of both species, on Brassica oleracea cabbage plants, in the presence or absence of a shared predator (Chrysoperla carnea larvae). Aphid population growth rates, aphid distributions on host plants and predator growth rates were measured. In single‐species treatments, C. carnea reduced M. persicae population growth rate, but had no significant impact on B. brassicae population growth rate, suggesting B. brassicae chemical defences are effective against C. carnea. Chrysoperla carnea had no significant impact on either aphid species population growth rate in mixed‐species treatments. Myzus persicae (ND) therefore experienced reduced predation in the presence of B. brassicae (CD) through a predator‐mediated indirect effect. Moreover, predator growth rates were significantly higher in the M. persicae‐only treatments than in either the B. brassicae‐only or mixed‐species treatments, suggesting predation was impaired in the presence of B. brassicae (CD). A trait‐mediated indirect interaction is proposed, consistent with associational resistance, in which the predator, upon incidental consumption of chemically‐defended aphids is deterred from feeding, releasing non‐defended aphids from predatory control.     

Differential effects of weather,plant phenology and predators on the seasonal variation of aphids on cabbage

The aphids Lipaphis erysimi pseudobrassicae (Davis) and Myzus persicae (Sulzer) pose serious threats to the production of cruciferous crops in the tropics. Understanding their population dynamics is important for developing integrated pest management programmes to minimize their damage to crops. This study investigated the effects of climatic factors, natural enemies and plant age on the population dynamics of these pests. The population density of aphids and their natural enemies in 20 cabbage plants, and weather conditions were monitored for five cropping seasons from 2019 to 2021 in two agroecological zones of Ghana (Coastal Savannah and Deciduous Forest zones). The highest population density of L. e pseudobrassicae was recorded in January (dry season) in both agroecological zones, while the highest population density for M. persicae occurred in September (minor rainy season) and August (dry spell) in the Coastal Savannah and Deciduous Forest zones, respectively. The highest aphid densities were noted to occur during periods with low relative humidity and low rainfall. The population density of L. e. pseudobrassicae was significantly negatively related to plant age, air temperature and relative humidity, and positively related to syrphids (Paragus borbonicus) and spiders in the Coastal Savannah zone, while in the Deciduous Forest zone, it was significantly positively related to coccinellids. On the other hand, M. persicae population density was significantly positively related to syrphids and coccinellids in the Deciduous Forest zone. Rainfall negatively affected syrphids in the Coastal Savannah zone, while air temperature positively affected syrphids and negatively affected spiders in the Deciduous Forest zone. Coccinellids had a significant positive relationship with relative humidity in the Deciduous Forest zone. This study provides important insights into the key factors that regulate aphid population densities on cabbage and will support development of timely interventions to manage these pests.     

Delineating the effects of a plant trait on interactions among associated insects

Plant traits can affect ecological interactions between plants, herbivores, and predators. Our study tests whether reduced leaf wax in peas alters the interaction between the pea aphid ( Acyrthosiphon pisum), a foliar-foraging predator (a lady beetle, Hippodamia convergens) and a ground-foraging predator (a ground beetle, Poecilus scitulus). We performed a 2×2×2 factorial experiment in which wax level, presence of H. convergens, and presence of P. scitulus were manipulated. Experimental arenas consisted of a cage surrounding three pea plants. One plant in each cage was stocked with 15 pea aphids. In greenhouse and field cage experiments, we assessed the effect of each factor and their interactions on aphid density. As in previous studies, H. convergens foraged for aphids more effectively on reduced wax peas than on normal peas. Other interactions among H. convergens, P. scitulus , and A. pisum were the same on both types of peas. We consider how aphid movement, plant growth, and a high frequency of predation by P. scitulus on H. convergens influenced pea aphid density.     

Effects of habitat fragmentation on the three-way interaction among ants,aphids and larvae of the giant purple emperor, <Emphasis Type="Italic">Sasakia charonda</Emphasis> (Hewitson), a near-threatened butterfly

Survival rates of both early and middle instar larvae of the nymphalid butterfly, Sasakia charonda, were estimated to be lowest on test trees planted in a meadow (site A), intermediate in a small, narrow secondary deciduous broadleaf forest (small patch, site B) and highest in a large secondary deciduous broadleaf forest (large forest, site C). The larval mortality rates due to predation by tree-climbing predators from the ground (tree climbing predator) such as ants and the larvae of carabids were estimated to be greater at sites A and B than those at site C. The number of predatory ants climbing test trees was significantly greater at sites A and B than at site C, and the ants harvested honeydew from aphids living on tree leaves at those two sites. Aphid densities were significantly higher on trees at sites A and B than at site C, and aphid densities and numbers of predatory ants were significantly and positively correlated at sites A and B. In an experiment controlling aphid density per branch on test trees, the numbers of ants and the mortality rates of S. charonda larvae were greater on branches with high aphid densities than on those with low aphid densities at both sites A and B. These results suggest that the aphid density per host tree was higher in the meadow and the small patch than in the large forest; at both sites these higher aphid densities attracted higher numbers of predatory ants to test trees, and as a result, mortality rates of S. charonda larvae were increased.     

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.     

Wedged between bottom-up and top-down processes: aphids on tansy

Abstract. 1. Many species of aphids exploit a single host‐plant species and have to cope with changing environmental conditions. They often vary greatly in abundance even when feeding on the same host. In a field experiment, the bottom‐up (plant quality/patch type frequency) and top‐down (ant attendance/predation) effects on the abundance of four species of aphids feeding on tansy (Tanacetum vulgare) were tested using a full factorial design. In addition, a model was used to examine these patch characteristics for their relative effects on the population dynamics and abundance of different aphid species. 2. Aphid numbers changed significantly depending on the quality of the host plant and the presence/absence of attending ants. The obligate myrmecophile, Metopeurum fuscoviride, was abundant on high‐quality plants, while on poor quality plants or on plants without attending ants these aphids did not survive until the end of the experiment. The facultative myrmecophiles, Aphis fabae and Brachycaudus cardui, and the unattended aphid species, Macrosiphoniella tanacetaria, all reached similar peak population densities, but M. tanacetaria did best in poor quality patches. 3. Natural enemies reduced aphid numbers, but those species feeding on high‐quality plants survived longer than those on poor‐quality plants, which existed only for a short period of time, especially when associated with ants. Losses due to migration of winged morphs and mortality caused by parasitoids were insignificant. 4. Varying the frequency of different patch types in a model indicates that different degrees of associations with ants are favoured in different environments. If the proportion of high‐quality patches in a habitat is large, obligate myrmecophiles do best. On increasing the number of poor‐quality patches, unattended species become more abundant. 5. The results suggest that, in spite of large species specific differences in growth rates, degree of myrmecophily or life cycle features, the temporal and spatial variability in top‐down and bottom‐up forces differentially affects aphid species and allows the simultaneous exploitation of a shared host‐plant species.     

Different-sized oak trees are equally protected by the aphid-tending ants

Theoretical models predict that the ecological impacts of ants on plants will vary with the size of the plant, but experimental evidence supporting this hypothesis is lacking. Focusing on aphid-tending ants, Lasius fuliginosus, and host oak trees, Quercus liaotungensis, we surveyed the density of aphids on the leaves of different-sized trees and the relationship between the densities of aphids and ants on the leaves. We then evaluated the impacts of ants on small (diameter at breast height (DBH) < 5 cm) and large trees (DBH > 25 cm). The field survey showed that the density of ants on leaves was determined by the density of aphids and the latter was similar among different-sized trees; through an ant-exclusion treatment, we found that the impacts of ants on plants, aphids, caterpillars, and galls did not differ by the size of the tree. L. fuliginosus had a significant protective effect for Q. liaotungensis, but this effect did not convert to enhanced fruit production during the 2-year study period. Considering that size can be used as an indicator of a plant’s ontogenetic stage, our results indicate that with similar densities of aphids and, in turn, ants on their canopy, trees at early and later ontogenetic stages are equally protected by aphid-tending ants.     

The role of ant-tended extrafloral nectaries in the protection and benefit of a Neotropical rainforest tree

One possible function of extrafloral nectaries is to attract insects, particularly ants, which defend plants from herbivores. We determined whether ants visiting saplings of the tree Stryphnodendronmicrostachyum (Leguminosae) provide protection (decreased plant damage due to ant molestation or killing of herbivores) and benefit (increased plant growth and reproduction associated with ant presence) to the plant. We compared ant and herbivore abundance, herbivore damage and growth of ant-visited plants and ant-excluded plants grown in sun and shade microhabitats of a 6-ha plantation in Costa Rica over a 7-month period. Results show that ants provided protection to plants not by reducing herbivore numbers but by molesting herbivores. Ants also reduced the incidence of pathogen attack on leaves. Protection was greater in the shade than in the sun, probably due to lower herbivore attack in the sun. Protection was also variable within sun and shade habitats, and this variability appeared to be related to variable ant visitation. Results also indicate that ant presence benefits the plant: ant-visited plants grew significantly more in height than ant-excluded plants. The cultivation of ants may serve as an important natural biological control in tropical forestry and agroforestry systems, where increased plant density can otherwise lead to increased herbivore attack. Received: 4 May 1998 / Accepted: 6 October 1998     

Mutualism in a community context: the positive feedback between an ant–aphid mutualism and a gall-making midge

Although mutualisms are widespread and often described in natural history accounts, their ecological influences on other community members remain largely unexplored. Many of these influences are likely a result of indirect effects. In this field study, we investigated the indirect effects of an ant–aphid mutualism on the abundance, survival rates and parasitism rates of a co-occurring herbivore. Rabdophaga salicisbrassicoides (Diptera: Cecidomyiidae) induces rosette galls on the developing shoots of Salix exigua trees, and populations can reach outbreak densities (up to 1,000 galls/stem) in central Washington State (USA). Ant-tended aphids feed on these same stems and often feed on gall tissue. In this study we used a combination of manipulative experiments and observational surveys to test the hypothesis that the abundances of aphids, ants, and galls have positive and reciprocal effects on one another, in a manner that would create a positive feedback loop in population growth. In addition, we examined whether the combined presence of ants and aphids reduces parasitism rates for the gallers. In support of the positive feedback loop hypothesis, aphids enjoyed higher population growth rates in the presence of ants and galls, the presence of ants and aphids resulted in increased abundance of galls, and the abundances of ants, aphids and galls were all positively correlated with one another. However, the mechanism underlying the positive effect of ants and aphids on galler density remains unknown, as the mutualism did not affect parasitism rates. More broadly, this study demonstrates that mutualisms can have significant and complex indirect effects on community and population ecology.     

Are ant-aphid associations a tritrophic interaction? Oleander aphids and Argentine ants

Summary Oleander aphids, (Aphis nerii), which are sporadically tended by ants, were used as a moded system to examine whether host plant factors associated with feeding site influenced the formation of ant-aphid associations. Seasonal patterns of host plant utilization and association with attendant ants were examined through bi-weekly censuses of the aphid population feeding on thirty ornamental oleander plands (Nerium oleander) in northern California in 1985 and 1986. Colonies occurred on both developing and senescing plant terminals, including leaf tips, floral structures, and pods. Aphids preferentially colonized leaf terminals early in the season, but showed no preference for feeding site during later periods. Argentine ants (Iridomyrmex humilis) occasionally tended aphid colonies. Colonies on floral tips were three to four times more likely to attract ants than colonies on leaf tips, even though the latter frequently contained more aphids. Ants showed a positive recruitment response to colonies on floral tips, with a significant correlation between colony size and number of ants. There was no recruitment response to colonies on leaf tips. These patterns were reproducible over two years despite large fluctuations in both aphid population density and ant activity. In a laboratory bioassay of aphid palatability, the generalist predator,Hippodamia convergens, took significantly more aphids reared on floral tips compared to those reared on leaf tips. The patterns reported here support the hypothesis that tritrophic factors may be important in modifying higher level arthropod mutualisms.     

Effects of Aphis fabae Scop, and of its attendant ants and insect predators on yields of field beans (Vicia faba L.)

Predators (mainly coccinellid adults and larvae and syrphid larvae), although few, were important in decreasing numbers of Aphis fabae on a small plot of field beans during the early stages of infestation in a year favourable to the aphid. At the same time, ants (Lasius niger L.), attending aphids on other plants on the same plot, effectively protected the aphids from predators for about 2 weeks, enabling the attended aphids to multiply faster than the unattended. When all aphid populations started to decline, predators became more numerous and accelerated the decline on both sets of plants. Bean plants without aphids yielded fifty-six seeds per plant; those with aphids but free from ants gave seventeen; and those with ant-attended aphids, eight seeds per plant. The damage and loss of yield was caused by the large aphid populations that developed when the pods were maturing, and not by the fewer aphids present when the plants were in flower. It appears that small, temporary infestations during flowering might increase the yield of field beans.     

A facultative mutualism between aphids and an invasive ant increases plant reproduction

1. The consequences to plants of ant–aphid mutualisms, particularly those involving invasive ants, are poorly studied. Ant–aphid mutualisms may increase or decrease plant fitness depending on the relative cost of herbivory by ant‐tended aphids versus the relative benefit of increased ant suppression of other (non‐aphid) herbivores. 2. We conducted field and greenhouse experiments in which we manipulated the presence and absence of cotton aphids (Aphis gossypii) on cotton plants to test the hypothesis that a mutualism between cotton aphids and an invasive ant, the red imported fire ant (Solenopsis invicta), benefits cotton plants by increasing fire ant suppression of caterpillars. We also manipulated caterpillar abundance to test whether the benefit of the mutualism varied with caterpillar density. 3. We found that more fire ants foraged on plants with cotton aphids than on plants without cotton aphids, which resulted in a significant reduction in caterpillar survival and caterpillar herbivory of leaves, flower buds, and bolls on plants with aphids. Consequently, cotton aphids indirectly increased cotton reproduction: plants with cotton aphids produced 16% more bolls, 25% more seeds, and 10% greater seedcotton mass than plants without aphids. The indirect benefit of cotton aphids, however, varied with caterpillar density: the number of bolls per plant at harvest was 32% greater on plants with aphids than on plants without aphids at high caterpillar density, versus just 3% greater at low caterpillar density. 4. Our results highlight the potential benefit to plants that host ant–hemipteran mutualisms and provide the first experimental evidence that the consequences to plants of an ant–aphid mutualism vary at different densities of non‐aphid herbivores.     

Community‐wide impact of an exotic aphid on introduced tall goldenrod

1. The aphid Uroleucon nigrotuberculatum Olive, which is specialised to the tall goldenrod, Solidago altissima L., in its native range, has become a dominant species on the introduced tall goldenrod in Japan. How this exotic aphid influenced arthropod communities on the introduced tall goldenrod in aphid‐present (spring) and aphid‐absent (autumn) seasons was examined, using an aphid removal experiment. 2. In spring, aphid presence increased ant abundance because aphid honeydew attracted foraging ant workers. A significant negative correlation was found between the numbers of ants and herbivorous insects other than aphids on the aphid‐exposed plants, but no significant correlation was detected on the aphid‐free plants. Thus, the aphid presence was likely to decrease the abundance of co‐occurring herbivorous insects through removal behaviour of the aphid‐tending ants. There were no significant differences in plant traits between the aphid‐exposed and aphid‐free plants. 3. In autumn, the numbers of lateral shoots and leaves, and the leaf nitrogen content were increased in response to the aphid infestation in spring. Because of the improvement of plant traits by aphid feeding, the abundance of leaf chewers increased on aphid‐exposed plants. In contrast, the abundance of sap feeders decreased on the aphid‐exposed plants. In particular, the dominant scale insect among sap feeders, Parasaissetia nigra Nietner, decreased, followed by a decrease in the abundance of ants attending P. nigra. Thus, aphid feeding may have attenuated the negative impacts of the tending ants on leaf chewers. 4. Aphid presence did not change herbivore species richness but changed the relative density of dominant herbivores, resulting in community‐wide effects on co‐occurring herbivores through ant‐mediated indirect effects, and on temporally separated herbivores through plant‐ and ant‐mediated indirect effects. The aphid also altered predator community composition by increasing and decreasing the relative abundance of aphid‐tending ants in the spring and autumn, respectively.     

Should aphids attract or repel ants? Effect of rival aphids and extrafloral nectaries on ant–aphid interactions

Among plants and herbivores, two types of conflicts occur in relation to mutualism with ants: one is competition for ant mutualism among myrmecophilous herbivores and plants, and the other is the conflict whether to attract or repel ants between myrmecophiles and nonmyrmecophiles that are damaged by ants. We investigated the extent to which two species of aphids (Megoura crassicauda and Aphis craccivora) and extrafloral nectaries on their host plant (Vicia faba var. minor) interact with one another for their relationships with ants. We designed an experiment where ants can choose to visit seedlings colonized by (1) M. crassicauda, (2) A. cracivora, (3) both aphid species, or (4) neither aphid species. Ants preferred A. craccivora to extrafloral nectaries and avoided tending M. crassicauda. We also analyzed the population growth of each aphid when it coexists with (1) ants, (2) the other aphid species, (3) ants and the other aphid species, or (4) neither of them. Under ant-free conditions, we detected an exploitative competition between the two aphid species. The ants had no significant effect on the population of A. craccivora, whereas they had negative effects on the population growth of M. crassicauda by attacking some individuals. When both aphids coexisted, M. crassicauda suffered ant attack more intensely because A. craccivora attracted more ants than extrafloral nectaries despite ant-repelling by M. crassicauda. On the other hand, the ants benefited A. craccivora by eliminating its competitor. To avoid ant attack, aphids may have been selected either to be more attractive to ants than other sympatric sugar sources or to repel the ants attracted to them. We hypothesize that competition among sympatric sugar sources including rival aphids and extrafloral nectaries is a factor restricting aphids to be myrmecophilous. Received: January 17, 2000 / Accepted: July 4, 2000