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Many ants and hemipterans are bound by a mutualistic relationship (trophobiosis) which represents an ecological keystone: ants receive food (honeydew) providing different kinds of protection. Partner protection against arthropod natural enemies is considered to be frequent and to interfere with biological control strategies of hemipteran pests.We carried out an ant‐exclusion field experiment in a fava bean (Vicia faba) plantation in Italy to monitor the abundance and behaviour of the ants (Plagiolepis pygmaea, Tetramorium semilaeve and the exotic Nylanderia jaegerskioeldi) and their relationship with facultative mutualist (Aphis fabae) and non‐mutualist (Megoura viciae) aphids, the arthropod natural enemies of the aphids, and extrafloral nectaries.Ants concentrated their activity on the attendance of facultative mutualist aphids much more than on the extrafloral nectaries. The ant‐exclusion treatment had no effect on the abundance of M. viciae and on the parasitization rate of A. fabae, while it reduced the abundance of A. fabae, aphid predators, and aphid parasitoids.Our results demonstrate that ant attendance does not imply the protection of aphids from arthropod natural enemies and suggest that the identity of the ant species involved is important for the outcome. As relatively few species have been studied in this regard, extending our knowledge to the role of more ant species is desirable to understand the ecology and evolution of ant‐aphid mutualisms and to refine integrated control strategies of aphid pests. [ABSTRACT FROM AUTHOR]

Bumblebees are opportunistic resource consumers and have been known to visit extrafloral nectaries and harvest aphid and scale honeydew. We report the first observation of bumblebees foraging psyllid honeydew in a tritrophic system in the boreal forest of Yukon Territory, Canada.In this system, a willow (Salix alaxensis) develops bizarrely enlarged petiole bases which act as nursery cavities for large numbers of psyllid immatures of Cacopsylla macleani (Homoptera: Psyllidae), with later instars dispersing over the underside of the leaf.These immatures produced abundant honeydew which was vigorously and persistently foraged by several bumblebees of multiple species (we identified Bombus vancouverensis, Bombus melanopygus and Bombus mckayi). Ants and vespid wasps were also present but in lower numbers.Salix alaxensis and Salix‐feeding psyllids are common throughout arctic and boreal regions, so honeydew production is potentially a supplementary food source for Bombus when floral nectar flows are limited due to low temperatures or drought. [ABSTRACT FROM AUTHOR]

The effects of drought‐induced changes in plant quality on aphid performance and population growth is well‐studied. The response of aphid behaviour to plant water limitation has received less attention. Water limitation may affect host‐plant colonization by altering the attractiveness of plants. Additionally, plant water limitation may inhibit feeding site establishment and phloem ingestion.Our goal was to examine bird cherry‐oat aphid (Rhopalosiphum padi L.) host selection and feeding behaviour under water limitation. We assessed aphid response to well‐watered, mildly‐stressed, and highly‐stressed wheat (Triticum aestivum L.) by evaluating (i) host‐plant selection through two‐choice assays, (ii) feeding behaviour using the electrical penetration graph technique, and (iii) phloem ingestion by quantifying honeydew production.Aphids were less likely to select highly stressed plants than a mildly stressed or well‐watered alternative. Aphids did not distinguish between mildly stressed and well‐watered plants. Aphid feeding behaviours, including duration of phloem ingestion, were not affected by water availability. However, honeydew production was reduced under both levels of water limitation. These results suggest that the volume of phloem ingested by aphids per unit time declined on stressed plants. The combination of lower colonization and diminished access to food on stressed plants may lead to a reduction in aphid abundance, independent of the direct effects of nutrition on individual aphid performance.This study highlights the potential contribution of herbivore behaviour to documented changes in aphid abundance on stressed plants and underscores the important role of plant water stress intensity in mediating plant‐herbivore interactions. [ABSTRACT FROM AUTHOR]

1. Mutualism between ants and honeydew‐excreting hemipterans is ubiquitous in the ecosystem. It is widely accepted that ant tending facilitates the colony growth of hemipterans by protecting them from predators and parasitoids. However, few studies have explored how ant tending helps defend against natural enemies. 2. Ghost ant Tapinoma melanocephalum and the invasive mealybug Phenacoccus solenopsis have a close mutual relationship. Previous studies have shown that ghost ant tending can definitely reduce parasitism and visit frequency of Aenasius bambawalei, the dominant endoparasitoid of P. solenopsis. However, the ghost ant workers seldom attack the parasitoids. It is still unclear how the ghost ant adversely affects parasitoids. This study explored the mechanism underlying the impacts of ants on natural enemies of the mealybugs. 3. Honeydew produced by P. solenopsis was an attractant to A. bambawalei. Parasitoids exhibited less searching activity, shorter longevity and lower parasitism when supplied with less honeydew. Aenasius bambawalei showed significant avoidance of pygidial gland secretions and visual cues of ghost ants. Parasitism in plants treated with 6‐methyl‐5‐hepten‐2‐one, actinidine, and gland extracts was significantly lower than that in plants treated only with solvents (paraffin oil or double‐distilled water). 4. It is concluded that honeydew consumption by ghost ants could negatively influence the performance of parasitoids. The pygidial gland secretions and visual cues of ghost ants also significantly inhibit the parasitism. These results may contribute to a better understanding of the regulation mechanism in ant–hemipteran–enemy interactions. [ABSTRACT FROM AUTHOR]

1. Uptake of environmental contaminants by lower trophic groups can have negative effects on higher trophic groups. This study tested the ability of selenium, an environmental contaminant found in high concentrations throughout the tissues of certain accumulating plants, to be transferred to ants via aphid tissue and honeydew. 2. Plants of the selenium accumulator, Raphanus sativus (wild radish), were watered with three different selenium treatments (0, 0.25 or 0.5 µg Se ml−1). Aphids, Myzus persicae, and Argentine ant colonies, Linepithema humile, were added to each caged plant and allowed to interact freely. Ant colonies were supplemented with one of three different food options to encourage the consumption of aphids, aphid honeydew, or aphids and honeydew. 3. The accumulation of selenium by each trophic group and a trophic transfer factor ( TTF) was calculated. The TTF for plants to aphids was > 1, indicating biomagnification, whereas the TTF for aphids to worker ants was < 1, indicating only biotransfer. Accumulated levels by worker ants did not statistically differ as a result of diet. 4. The amounts of selenium acquired by ants as a factor of diet and caste were compared. Plants, aphids and worker ants accumulated selenium in a dose-dependent manner. Ant queens did not contain detectable amounts of selenium. Honeydew contained comparable amounts of selenium to plant selenium levels. 5. Access to toxic compounds via honeydew and insect protein may have negative effects on the range expansion of invasive species, such as the Argentine ant. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

1. Ants, as well as many species of parasitoids and predators, rely on sugar-rich foods such as honeydew to fulfill their energetic needs. Thus, ants and natural enemies may interact through the shared honeydew exploitation. 2. Ant-exclusion experiments were performed in a citrus orchard to test the hypothesis that ants may impact the energy reserves of predators and parasitoids through the competition for honeydew. Through the use of high-performance liquid chromatography ( HPLC) the level of ant activity with the energy reserves and feeding history of individual specimens collected in the field during representative days of spring, summer, and autumn were related. 3. Out of 145 Aphytis chrysomphali Mercet parasitoids captured in the field, 65% were classified as sugar-fed and 24.7% as honeydew-fed. In summer, when ant activity peaked, there was a significant negative correlation between the level of ant activity and the total sugar content and honeydew feeding incidence by A. chrysomphali. Out of 47 individuals of the predator Chrysoperla carnea sensu lato (Stephens), captured in the field, 55.3% were classified as sugar-fed. We found a significant negative effect of the level of ant activity on the sugar-feeding incidence by C. carnea in spring. 4. The present study provides evidence that ants can interfere with the energy reserves of natural enemies. This interaction may be widespread in various ecosystems with important consequences for the arthropod community composition and with practical implications for biological control given that absence of sugar feeding is detrimental for the fitness of many species of predators and parasitoids. [ABSTRACT FROM AUTHOR]

1. Plants take nutrients from the rhizosphere via two pathways: (i) by absorbing soil nutrients directly via their roots and (ii) indirectly via symbiotic associations with nutrient-providing microbes. Herbivorous insects can alter these pathways by herbivory, adding their excrement to the soil, and affecting plant-microbe associations. 2. Little is known, however, about the effects of herbivorous insects on plant nutrient uptake. Greenhouse experiments with soybean, aphids, and rhizobia were carried out to examine the effects of aphids on plant nutrient uptake. 3. First, the inorganic soil nitrogen and the sugar in aphid honeydew between aphid-infected and -free plants were compared. It was found that aphid honeydew added 41 g m−2 of sugar to the soil, and that aphids decreased the inorganic soil nitrogen by 86%. This decrease may have been caused by microbial immobilisation of soil nitrogen followed by increased microbial abundance as a result of aphid honeydew. 4. Second, nitrogen forms in xylem sap between aphid-infected and -free plants were compared to examine nitrogen uptake. Aphids decreased the nitrogen uptake via both pathways, and strength of the impact on direct uptake via plant roots was greater than indirect uptake via rhizobia. The reduced nitrogen uptake by the direct pathway was as a result of microbial immobilisation, and that by the indirect pathway was probably because of the interaction of microbial immobilisation and carbon stress, which was caused by aphid infection. 5. The present results demonstrate that herbivorous insects can negatively influence the two pathways of plant nutrient uptake and alter their relative importance. [ABSTRACT FROM AUTHOR]

Protection mutualisms between ants and honeydew-producing insects influence arthropod food webs by altering the behaviour, activity and local abundance of ants on plants. Ants often tend multiple species of honeydew-producing insects; however, studies that examine such effects typically consider only pairwise mutualisms. This study investigates how multi-species mutualisms between ants and honeydew-producing insects structure arthropod food webs in citrus., In an organic lemon orchard in San Diego County, California, U.S.A., ants or honeydew-producing insects (or neither) were experimentally removed from individual, mature lemon trees and then abundances of the following were estimated over a 2-year period: the Argentine ant, five species of honeydew-producing hemipterans, and California red scale. Red scale produces no honeydew but indirectly benefits from the presence of ants, which disrupt parasitism by Aphytis wasps., Mutualism between ants and honeydew-producing hemipterans indirectly and positively affected red scale. Levels of parasitism decreased with increasing ant recruitment, and red scale density increased with both increasing ant recruitment and increasing honeydew-producing hemipteran abundance. Moreover, abundances of each of three honeydew-producing hemipteran species emerged as positive predictors of red scale density in different analyses; this finding suggests that individual hemipteran taxa exhibit distinct and spatially localised effects on ants., Evidence of positive, indirect effects between two focal species of honeydew-producing Hemiptera was also detected. Guilds of honeydew-producing hemipterans may provide ants with honeydew more consistently or for longer than that produced by any single species. These results argue for an appreciation of how mutualist diversity affects the ecological consequences of mutualisms. [ABSTRACT FROM AUTHOR]

Adult parasitoids depend on sugar-rich foods such as nectar and honeydew to meet their energy requirements and control insect pests. However, it is poorly known whether parasitoids can detect and feed on honeydew in agroecosystems, where it is the primary carbohydrate source, because this sugar source is less apparent in comparison to nectar and sometimes contains repellent compounds for parasitoids., High-performance liquid chromatography ( HPLC) analyses were carried out to test whether Aphytis melinus DeBach (Hymenoptera: Aphelinidae), a parasitoid whose host does not produce honeydew, feeds on honeydew from non-hosts. In addition, the correlation between the parasitoid's sugar reserves and honeydew abundance was determined. To do this, both the levels of honeydew producers and the sugar levels of individual collected parasitoids were assessed during different seasons. The overall sugar content was treated as an indicator of energy reserves and the erlose-melezitose ratio as an indicator of honeydew feeding., The data show that A. melinus fed commonly on honeydew from non-host hemipterans. More than 50% of the female parasitoids collected in spring and summer had recently fed on honeydew and most of them showed a high sugar content., However, in autumn, when the number of honeydew producers was three times lower than in spring and summer, less than 20% of A. melinus were found to have fed on honeydew, with the average total sugar content being reduced by a factor of three., This study demonstrates that A. melinus commonly feeds on honeydew in the field, even though its host does not produce honeydew. The results also suggest that the exploitation of honeydew by A. melinus is a function of the density and species of honeydew producers. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

1. Environmental cues associated with prey are known to increase predator foraging efficiency. Ladybird larvae are major predators of aphids. The sugary excretion of aphids (honeydew) has been proposed to serve as a prey-associated cue for ladybird larvae. 2. Ladybird larvae are regularly found on the ground moving between plants or after falling off plants. The use of prey-associated cues would be particularly beneficial for ladybird larvae on the ground in that such cues would help them to decide which plants to climb because aphids are patchily distributed within as well as amongst plants and, as a result, many plants are either not infested with aphids or do not host an aphid species of high nutritional value for ladybird larvae. 3. Laboratory experiments with larvae of Hippodamia convergens Guérin-Méneville (Coleoptera: Coccinellidae) were carried out to explore whether honeydew accumulated on the ground is used as a foraging cue. The study also investigated whether, if honeydew is a foraging cue, larvae show differential responses to honeydew of high-quality prey Acyrthosiphon pisum Harris compared with that of low-quality prey Aphis fabae Scopoli (both: Homoptera: Aphididae). 4. Hippodamia convergens larvae stayed longer in areas containing honeydew but did not engage in longer bouts of searching. Furthermore, larvae did not distinguish between honeydew from high- and low-quality aphid prey. [ABSTRACT FROM AUTHOR]

1. To maximise their reproductive success, the females of most parasitoids must not only forage for hosts but must also find suitable food sources. These may be nectar and pollen from plants, heamolymph from hosts and/or honeydew from homopterous insects such as aphids. 2. Under laboratory conditions, females of Cotesia vestalis, a larval parasitoid of the diamondback moth ( Plutella xylostella) which does not feed on host blood, survived significantly longer when held with cruciferous plants infested with non-host green peach aphids ( Myzus persicae) than when held with only uninfested plants. 3. Naïve parasitoids exhibited no preference between aphid-infested and uninfested plants in a dual-choice test, but those that had been previously fed aphid honeydew significantly preferred aphid-infested plants to uninfested ones. 4. These results suggest that parasitoids that do not use aphids as hosts have the potential ability to learn cues from aphid-infested plants when foraging for food. This flexible foraging behaviour could allow them to increase their lifetime reproductive success. [ABSTRACT FROM AUTHOR]

1. We monitored three different-sized wood ant ( Formica aquilonia Yarrow) mounds over a 3-year period in Finnish boreal forests dominated by Norway spruce ( Picea abies Karst.), to assess the seasonal temperature dependency of ant activity. Additionally, we also monitored Norway spruce trees around the mounds for descending honeydew foragers. 2. The amount of collected honeydew and prey and its composition, as well as the carbon (C), nitrogen (N), and phosphorus (P) in honeydew and invertebrate prey was also investigated. 3. The number of warm days (average temperature above 20 °C) and the amount of precipitation differed among the years. Ant activity at the mounds (but not on the trees) was highly correlated with air temperature throughout the ant-active season (May–September), but ant activity in spring and autumn was lower than in summer at similar temperatures. During all 3 years, honeydew played a major role in wood ant nutrition (78–92% of dry mass). Invertebrate prey was mainly Diptera (on average 26.2%), Coleoptera (12.5%), Aphidina (9.3%), and Arachnoida (8.5%). 4. The total amounts of C, N, and P input brought into the ant mounds in the form of food (both honeydew and prey) on the stand level were 12.6–39.0, 1.6–4.6 and 0.1–0.4 kg ha−1 year−1, respectively, which is equivalent to 2–6%, 12–33% and 27–58% of the fluxes in annual needle litterfall in typical boreal Norway spruce forests. Thus, wood ants can play a significant role in short term and local N and P cycling of boreal forest ecosystems. [ABSTRACT FROM AUTHOR]

1. The use of flowering vegetation has been widely advocated as a strategy for providing parasitoids and predators with nectar and pollen. However, their herbivorous hosts and prey may exploit floral food sources as well. 2. Previous laboratory studies have shown that not all flower species are equally suitable in providing accessible nectar. Relatively little is known about actual nectar exploitation under field conditions. 3. The present study investigates nectar exploitation by the pest, Plutella xylostella, and its parasitoid, Diadegma semiclausum, under field conditions and examines whether floral nectar exploitation in the field can be predicted based on controlled laboratory studies. 4. Insects were collected from fields bordered by flowering margins containing Fagopyrum esculentum, Lobularia maritima, Anethum graveolens, Centaurea jacea or the grass Lolium perenne (control). Whole insect bodies were individually assayed by HPLC to establish their sugar profile as a measure of the level of energy reserves and the degree of food source use. 5. The average overall sugar content of P. xylostella and D. semiclausum collected in fields bordered by flowering margins was significantly higher than those of individuals collected from grass-bordered control plots. To the authors’ knowledge, this represents the first demonstration that nectar-providing plants enhance the energetic state of herbivores under field conditions. 6. In contrast to earlier laboratory studies, the present study did not find elevated sugar contents in P. xylostella and D. semiclausum individuals collected from fields bordered by buckwheat ( F. esculentum). 7. The present study shows widespread sugar feeding by both the herbivore and its parasitoid. It also shows that laboratory studies establishing nectar exploitation under controlled conditions can not always be extrapolated to actual exploitation under field conditions. This emphasises the importance of studying field-collected insects with regard to food source use and nutritional status. [ABSTRACT FROM AUTHOR]

1. Although interactions between ants and honeydew-producing insects have received considerable study, relatively little is known about how these interactions alter the behaviour of ants in ways that affect other arthropods. In this study, field and greenhouse experiments were performed that examined how the presence of aphids ( Aphis fabae solanella) on Solanum nigrum influenced the foraging behaviour of Argentine ants ( Linepithema humile) and, in turn, modified the extent to which ants deter larval lacewings ( Chrysoperla rufilabris), which are known aphid predators. 2. A field experiment demonstrated that the level of foliar foraging by ants increased linearly with aphid abundance, whereas no relationship existed between the level of ground foraging by ants and aphid abundance. 3. In the greenhouse, as in the field, foliar foraging by ants greatly increased when aphids were present. Higher levels of foliar foraging led to a twofold increase in the likelihood that ants contacted aphid predators. As a result of these increased encounters with ants, lacewing larvae were twice as likely to be removed from plants with aphids compared with plants without aphids. Once contact was made, however, the behaviour of ants towards lacewing larvae appeared similar between the two experimental groups. 4. Argentine ants drive away or prey upon a diversity of arthropod predators and parasitoids, but they also exhibit aggression towards certain herbivores. Future work should attempt to quantify how the ecological effects that result from interactions between honeydew-producing insects and invasive ants, such as L. humile, differ from those that result from interactions between honeydew-producing insects and native ants. [ABSTRACT FROM AUTHOR]

1. The Argentine ant, Linepithema humile, tends honeydew-excreting homopterans and can disrupt the activity of their natural enemies. This mutualism is often cited for increases in homopteran densities; however, the ant’s impact on natural enemies may be only one of several effects of ant tending that alters insect densities. To test for the variable impacts of ants, mealybug and natural enemy densities were monitored on ant-tended and ant-excluded vines in two California vineyard regions. 2. Ant tending increased densities of the obscure mealybug, Pseudococcus viburni, and lowered densities of its encyrtid parasitoids Pseudaphycus flavidulus and Leptomastix epona. Differences in parasitoid recovery rates suggest that P. flavidulus was better able to forage on ant-tended vines than L. epona. 3. Densities of a coccinellid predator, Cryptolaemus montrouzieri, were higher on ant-tended vines, where there were more mealybugs. Together with behavioural observations, the results showed that this predator can forage in patches of ant-tended mealybugs, and that it effectively mimics mealybugs to avoid disturbance by ants. 4. Ant tending increased densities of the grape mealybug, Pseudococcus maritimus, by increasing the number of surviving first-instar mealybugs. Parasitoids were nearly absent from the vineyard infested with P. maritimus. Therefore, ants improved either mealybug habitat or fitness. 5. There was no difference in mealybug distribution or seasonal development patterns on ant-tended and ant-excluded vines, indicating that ants did not move mealybugs to better feeding locations or create a spatial refuge from natural enemies. 6. Results showed that while Argentine ants were clearly associated with increased mealybug densities, it is not a simple matter of disrupting natural enemies. Instead, ant tending includes benefits independent of the effect on natural enemies. Moreover, the effects on different natural enemy species varied, as some species thrive in the presence of ants. [ABSTRACT FROM AUTHOR]

1. To clarify the use of honeydew in foraging for aphids by larvae of the ladybird beetle, Coccinella septempunctata L., searching behaviour of ladybird larvae for Aphis craccivora Koch and Acyrthosiphon pisum Harris and the abundance of honeydew under aphid colonies were examined in laboratory experiments. 2. More larvae responded by climbing the plants with aphids than responded to plants without aphids. When the plants were replaced with sticks, in order to exclude visual and olfactory cues from plants and aphids, more larvae of C. septempunctata climbed sticks above the area that contained honeydew than climbed sticks above the area that did not contain honeydew. Then, ladybird larvae use honeydew as a contact kairomone when foraging for aphids. 3. Aphis craccivora deposited a larger number of honeydew droplets beneath the plants than did similar numbers of A. pisum. Thus, C. septempunctata larvae licked more frequently the honeydew of A. craccivora than that of A. pisum and spent longer searching on the area containing honeydew of A. craccivora than that of A. pisum. Consequently, a larger number of larvae climbed a stick above honeydew of A. craccivora than that of A. pisum. 4. It may be also considered that C. septempunctata larvae can distinguish honeydew of the two aphid species and respond more strongly to A. craccivora than A. pisum. [ABSTRACT FROM AUTHOR]

1. The degree of infestation by New Zealand sooty beech scale insects ( Ultracoelostoma assimile, Homoptera: Margarodidae) varies dramatically among adjacent southern beech trees ( Nothofagus spp., Fagaceae), but has previously been assumed to be uniformly or randomly distributed within individual host trees. In this study, a full-census survey was conducted from ground level to canopy level on 14 naturally occurring, canopy-dominant red beech ( Nothofagus fusca) trees (size range 38.7–107.6 cm diameter at breast height) to determine the degree of within-tree heterogeneity in herbivore density. 2. The within-tree distribution of the sooty beech scale was vertically stratified and highly heterogeneous, with the greatest densities occurring on bark surfaces in the canopy rather than on the trunk, and on the lower rather than upper sides of the branches. The spatial distribution was strongly negatively correlated with trunk and branch diameter, and increasing bark thickness (as a function of diameter) provides a plausible explanation for differences in the establishment and population density of sooty beech scale insects with trunk and branch size. Furthermore, there was a significant change in the spatial distribution of scale insect populations on trunks and branches of trees of increasing diameter at breast height. This indicates a strong temporal component to the spatial dynamics of the sooty beech scale insect driven by changing host phenology. Future studies on phytophagous insects infesting large host trees need to consider more explicitly changes in population dynamics through space and time. 3. Because of the high degree of within-tree heterogeneity in population density, the total population size of scale insects on an individual tree could not be predicted from any measure of population density low on the trunk. However, the dry weight biomass of sooty mould fungi growing on the ground beneath infested trees was a remarkably accurate predictor of the total population size of scale insects. The use of sooty mould fungi as a relative measure of population size could be incorporated into studies of other honeydew-producing hemipterans, since the growth of sooty mould is a distinctive feature synonymous with high concentrations of honeydew production worldwide. [ABSTRACT FROM AUTHOR]

1. Intraspecific competition can be an important factor influencing the individual fitness of organisms. This study reports density-dependent effects on the fecundity of the beech scale insect ( Ultracoelostoma assimile, Hemiptera: Margarodidae) on naturally occurring, canopy-dominant red beech ( Nothofagus fusca, Fagaceae) trees in New Zealand. For the first time an increasing intensity of intraspecific competition at increasing spatial scales within individual host trees is demonstrated. 2. Beech scale insect ‘tests’ containing adult female scale insects and eggs were collected from the trunks of 10 red beech trees [17.7–48.5 cm in diameter at breast height (1.4 m), d.b.h.] with varying densities of scale insect infestation. The relationship between individual female fecundity and scale insect density at three spatial scales: local (within 5 cm), lower trunk (below 2 m above ground level), and whole tree, was tested. 3. Beech scale insect fecundity was density dependent, with total female egg load decreasing with increasing scale insect density. The strength of the density-dependent effect increased with increasing spatial scale, suggesting that scale insects are creating a tree-wide drain on the quality of phloem sap, rather than depleting nutrients from localised, high-density areas within trees. These results indicate that at high densities, the New Zealand beech scale insect can have a negative effect on the nutritional quality of the phloem of red beech, thus negatively affecting conspecifics elsewhere on individual host trees. The increasing intensity of the effect with increasing spatial scale within individual trees emphasises the importance of measuring density-dependent effects at the appropriate spatial scale. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Summary1. The size–distance relationship among honeydew-collecting foragers of the red wood ant Formica rufa was investigated. Within the colony territory, the size (as measured by head width) and fresh weight of samples of foragers were determined for ants ascending and descending trees near, and farther from, the central nest mound. 2. The mean size of the ants was significantly higher at far trees than at near trees in six out of the seven colonies investigated, confirming the general presence of the size–distance relationship. 3. In three colonies, a load–distance relationship was also found. For a given head width, honeydew-carrying ants descending far trees were significantly heavier than those descending near trees (i.e. they were carrying heavier loads from trees farther away from the central nest mound). 4. This is the first time that both load–distance and size–distance relationships have been reported in foraging workers from the same ant colony. 5. The combined effects of these characteristics suggest that colony foraging efficiency is enhanced by far trees being visited by the larger workers that then return with heavier loads of honeydew. [ABSTRACT FROM AUTHOR]

Abstract. 1. The wasp Diaeretiella rapae uses honeydew emitted by its host, the cabbage aphid Brevicoryne brassica, as a kairomone (chemicals emitted by an organism as part of its activity and used by its natural enemies to their advantage). The role of the kairomone in foraging decisions by the parasitic wasp was explored by manipulating the amount of honeydew and the number of aphids in a colony independently. The count-down patch-exploitation mechanism (Iwasa et al., 1981) was employed to predict the results of these manipulations and contrast them with the predictions of Waage's (1979) model. 2. Kairomonal activity of honeydew decreased as the honeydew aged and lost its activity completely within 72 h. 3. The wasp was exposed to different amounts of honeydew on (a) aphid-free leaves and (b) leaves bearing colonies of 150 aphids. The parasitoid search time on both leaf types increased with increasing honeydew contamination. On aphid-bearing leaves, the number of attacked aphids in the colonies also increased with increasing honeydew contamination. The presence of aphids reduced the parasitoid search time compared to search time on aphid-bearing leaves with the same level of honeydew contamination, as predicted by the count-down model. 4. Parasitoids exposed to the same amount of honeydew, but a different number of aphids in a colony on the leal, first increased the number of aphids attacked in relation to the number of aphids in the colony. But as the colony reached a certain size, the number of aphids attacked levelled off, despite the increase in the number of aphids in the colony. Search time was variable but did not exhibit any trend as the number of aphids in the colonies increased. 5. These results suggest that honeydew level is used by D. rapae as a cue for assessment of the number of aphids in the colony. In such cases, a count-down exploitation mechanism gives the best results to a forager. Many parasitoids may use kairomones for patch assessment... [ABSTRACT FROM AUTHOR]