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Pest control in agriculture is mainly based on the application of insecticides, which may impact nontarget beneficial organisms leading to undesirable ecological effects. Neonicotinoids are among the most widely used insecticides. However, they have important negative side effects, especially for pollinators and other beneficial insects feeding on nectar. Here, we identify a more accessible exposure route: Neonicotinoids reach and kill beneficial insects that feed on the most abundant carbohydrate source for insects in agroecosystems, honeydew. Honeydew is the excretion product of phloem-feeding hemipteran insects such as aphids, mealybugs, whiteflies, and psyllids. We allowed parasitic wasps and pollinating hoverflies to feed on honeydew from hemipterans feeding on trees treated with thiamethoxam or imidacloprid, the most commonly used neonicotinoids. LC-MS/MS analyses demonstrated that both neonicotinoids were present in honeydew. Honeydew with thiamethoxam was highly toxic to both species of beneficial insects, and honeydew with imidacloprid was moderately toxic to hoverflies. Collectively, our data provide strong evidence for honeydew as a route of insecticide exposure that may cause acute or chronic deleterious effects on nontarget organisms. This route should be considered in future environmental risk assessments of neonicotinoid applications. [ABSTRACT FROM AUTHOR]

Bird exclusion experiments consistently show that birds exhibit strong top-down control of arthropods, including ants and the honeydew-producing insects (HPIs) that they tend. However, it remains unclear whether the results of these small-scale bird exclosure experiments can be extrapolated to larger spatial scales. In this study, we use a natural bird removal experiment to compare the prevalence of ants and HPIs between Guam, an island whose bird community has been extirpated since the 1980s due to the introduction of the brown tree snake, and two nearby islands (Rota and Saipan) that have more intact bird assemblages. Consistent with smaller-scale bird exclosure experiments, we show that (1) forest trees from Guam are significantly more likely to host HPIs than trees from Saipan and (2) ants are nearly four times as abundant on Guam than on both Saipan and Rota. The prevalence of HPIs varied slightly based on tree species identity, although these effects were not as strong as island-level effects associated with bird loss. Ant community composition differed between Guam and the other two islands. These results corroborate past observational studies showing increased spider densities on Guam and suggest that trophic changes associated with landscape-level bird extirpation may also involve alterations in the abundance of ants and HPIs. This study also provides a clear example of the strong indirect effects that invasive species can have on natural food webs. [ABSTRACT FROM AUTHOR]

Honeydew production by Hemiptera is an ecologically important process that facilitates mutualisms and increases nutrient cycling. Accurate estimates of the amount of honeydew available in a system are essential for quantifying food web dynamics, energy flow, and the potential growth of sooty mould that inhibits plant growth. Despite the importance of honeydew, there is no standardized method to estimate its production when intensive laboratory testing is not feasible. We developed two new models to predict honeydew production, one based on insect body mass and taxonomic family, and one based on body mass and life stage. We tested the accuracy of both models’ predictions for a diverse range of honeydew-producing hemipteran families (Aphididae, Pseudococcidae, Coccidae, Psyllidae, Aleyrodidae, Delphacidae, Cicadellidae). The method based on body mass and family provided more accurate estimates of honeydew production, due to large variation in honeydew production among families. We apply our methodology to a case study, the recalculation of honeydew available to invasive red imported fire ant (Solenopsis invicta) in the United States. We find that the amount of honeydew may be an order of magnitude lower than that previously estimated (2.16 versus 21.6 grams of honeydew per day) and discuss possible reasons for the difference. We anticipate that being able to estimate honeydew production based on minimal biological information will have applications to agriculture, invasion biology, forestry, and carbon farming. [ABSTRACT FROM AUTHOR]

Honey bees, Apis species, obtain carbohydrates from nectar and honeydew. These resources are ripened into honey in wax cells that are capped for long-term storage. These stores are used to overcome dearth periods when foraging is not possible. Despite the economic and ecological importance of honey, little is known about the processes of its production by workers. Here, we monitored the usage of storage cells and the ripening process of honey in free-flying A. mellifera colonies. We provided the colonies with solutions of different sugar concentrations to reflect the natural influx of nectar with varying quality. Since the amount of carbohydrates in a solution affects its density, we used computer tomography to measure the sugar concentration of cell content over time. The data show the occurrence of two cohorts of cells with different provisioning and ripening dynamics. The relocation of the content of many cells before final storage was part of the ripening process, because sugar concentration of the content removed was lower than that of content deposited. The results confirm the mixing of solutions of different concentrations in cells and show that honey is an inhomogeneous matrix. The last stage of ripening occurred when cell capping had already started, indicating a race against water absorption. The storage and ripening processes as well as resource use were context dependent because their dynamics changed with sugar concentration of the food. Our results support hypotheses regarding honey production proposed in earlier studies and provide new insights into the mechanisms involved. [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]

In many ecosystems food-web dynamics are driven by spatial and temporal variation in the availability of sugar resources, which form the primary or even exclusive dietary constituents for many species. Scale insects (Hemiptera) produce sugar-rich honeydew, which can be a keystone sugar source in honeydew ecosystems worldwide. In New Zealand, most previous research in honeydew ecosystems has been conducted in areas where herpetofauna are heavily suppressed by introduced predators. Consequently, little is known about potential trophic interactions between endemic lizards and scale insects. Korapuki Island is one of the few remaining locations in New Zealand where endemic scale insects and lizards survive in densities likely to be representative of prehuman conditions. We examined the relative importance of different sugar resources on Korapuki Island to Duvaucel's geckos (Hoplodactylus duvaucelii) and common geckos (Woodworthia maculatus). We recorded the abundance and morphometrics of geckos attending five sugar-producing plant species (two of which host honeydew-producing scale insects) three times daily along a fixed transect. Large numbers of Duvaucel's and common geckos were recorded nocturnally feeding on honeydew produced by the scale insect Coelostomidia zealandica (Coelostomidiidae). Duvaucel's geckos of all sizes and genders fed extensively on honeydew throughout the year, favouring ngaio (Myoporum laetum) trees with high scale insect infestations, but were seldom recorded at other sugar resources. In contrast, juvenile common geckos were infrequently recorded on honeydew-producing trees. Common geckos fed on a variety of other sugar resources, with all sizes and sexes abundant on nectar and sap of flax (Phormium tenax) and seasonally exploiting nectar of pohutukawa (Metrosideros excelsa). The strength of interactions between scale insects and geckos, particularly for the Duvaucel's gecko population on Korapuki Island, indicates the importance of honeydew in addition to more ephemeral sugar resources such as nectar. Accordingly, the re-establishment of honeydew-producing Hemiptera populations should be considered in future conservation and restoration plans. [ABSTRACT FROM AUTHOR]

While many studies have demonstrated that ants provide beneficial services to aphids, Bristow (Ant-plant interactions, Oxford University Press, Oxford, 104-119, ) first questioned why so few aphid species are ant-attended. Phylogenetic trees have demonstrated multiple gains and loss of ant-attendance in the course of aphid-ant interactions, implying that mutualisms easily form and dissolve. Several studies have reported the factors that influence the formation and maintenance of aphid-ant interactions. Examples include the physiological costs of ant attendance, competition for mutualistic ants, ant predation on aphids, the influence of host plants, and parasitoid wasps. Recent physiological techniques have also revealed the chemical component of aphid-ant mutualisms. The honeydew of ant-attended aphids contains melezitose (a trisaccharide), which has an important role in aphid-ant interactions. Studies of cuticular hydrocarbons on aphids and ants have clarified the underlying mechanisms of ant predation on aphids. Attending ants also reduce aphid dispersal ability, causing the formation of fragmented aphid populations with low genetic diversity in each population. The reduced aphid dispersal could be partly explained by higher wing loading and reduction of flight apparatus due to ant attendance. Whether ant attendance is associated with the range of host plants of aphids or genetic variation in microorganism in aphids remain to be explored. [ABSTRACT FROM AUTHOR]

Honeydew produced by sooty beech scale insects ( Ultracoelostoma spp., Homoptera: Coelostomidiidae) is a keystone ecological process in New Zealand beech ( Nothofagus spp., Nothofagaceae) forest. This work puts forward a model of honeydew production based on individual insects that presumes feeding and excretion are episodic processes driven by the insect rather than the passive processes that were previously assumed. The model is parameterized using existing data and then compared to an independent pre-existing dataset. The model suggests that over a 12-h period, on average the insects suck sap for 2 h, and excrete waste sap for 12 min. Resource uptake by the insects appears to be limited by the time required to process the sap, consistent with the observed relationship between honeydew production rates and ambient temperature. This implies that insect feeding rates may be ultimately limited by the low nitrogen content of phloem sap. [ABSTRACT FROM AUTHOR]

The article discusses a study concerning the ant mimicking behavior of female aphid parasitoid, Lysiphlebus fabarum against black bean aphid, Aphis fabae. It is stated that females encountered an average of 47.4 aphids per hour and reduced their defensive behavior by using ant like antennation. It further states that the mimicking behavior of parasitoid females increased in food deprived condition which revealed a possibility of mutualism between parasitoid and the ant species.

Plant-ants live in a mutualistic association with host plants known as ''myrmecophytes'' that provide them with a nesting place and sometimes with extra-floral nectar (EFN) and/or food bodies (FBs); the ants can also attend sap-sucking Hemiptera for their honeydew. In return, plant-ants, like most other arboreal ants, protect their host plants from defoliators. To satisfy their nitrogen requirements, however, some have optimized their ability to capture prey in the restricted environment represented by the crowns of trees by using elaborate hunting techniques. In this study, we investigated the predatory behavior of the ant Azteca andreae which is associated with the myrmecophyte Cecropia obtusa. We noted that up to 8350 ant workers per tree hide side-by-side beneath the leaf margins of their host plant with their mandibles open, waiting for insects to alight. The latter are immediately seized by their extremities, and then spread-eagled; nestmates are recruited to help stretch, carve up and transport prey. This group ambush hunting technique is particularly effective when the underside of the leaves is downy, as is the case for C. obtusa. In this case, the hook-shaped claws of the A. andreae workers and the velvet-like structure of the underside of the leaves combine to act like natural Velcro® that is reinforced by the group ambush strategy of the workers, allowing them to capture prey of up to 13,350 times the mean weight of a single worker. [ABSTRACT FROM AUTHOR]

Marchalina hellenica is the main honeydew producing insect of pine trees. It is endemic to Greece and Turkey and introduced to the Italian island of Ischia. It has one generation per year and the adult females appear on the trees only after mid March. Studies on the genetic structure of the insect show that the low genetic variability may be due to the fact that it can not be dispersed long distances in correlation with the parthenogenetic reproduction. The amount of honeydew produced by the insect varies over the year and mainly depends on the size and age of the nymphs. [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]

Trophic interactions involving predators, herbivores, and plants have been described in terrestrial systems. However, there is almost no information on the effect of trophic interactions on microbial phyllosphere community abundance, diversity, or structure. In this study, the interaction between a parasitoid, an insect herbivore, and the fungal phyllosphere community is examined. Parasitoid wasps have an indirect negative impact on fungal community diversity. On the citrus phyllosphere, the exotic wasp species, Amitus hesperidum and Encarsia opulenta, may parasitize the citrus blackfly ( Aleurocanthus woglumi). If parasitism levels are low, the blackfly may produce significant amounts of honeydew secretions on the surface of the leaf. Honeydew deposition provides a carbon-rich substrate for the development of fungal growth persisting as sooty mold on the leaves. Leaves from sooty mold-infested grapefruit ( Citrus paradisi) trees were collected from multiple orchards in south Texas. The effect of different levels of exotic parasite activity, citrus blackfly, and sooty mold infestation on phyllosphere mycobiota community structure and diversity was examined. Our results suggest the presence of the parasitoid may lead to a top–down trophic cascade affecting phyllosphere fungal community diversity and structure. Additionally, persistent sooty mold deposits that have classically been referred to as Capnodium citri (and related asexual morphological forms) actually comprise a myriad of fungal species including many saprophytes and potential fruit and foliar pathogens of citrus. [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]

Sugar-rich honeydew production by phloem-feeding sooty beech scale insects (Ultracoelostoma spp.) is a keystone ecological process in New Zealand beech (Nothofagus) forest. A dynamical model to describe the formation of individual honeydew droplets is derived using simple fluid dynamics. The model assumes that the insects play little part in regulating flow, and that flow is driven by phloem hydrostatic pressure and regulated by environmental conditions affecting water evaporation from droplets. The model predicts different types of behaviour depending on local environmental conditions. In dry conditions, evaporation rates are high and honeydew droplets can become so concentrated that further excretion is prevented and droplet formation ceases. In humid conditions, the high concentrations required to prevent droplet formation do not occur and droplets continue to grow indefinitely. The model is fitted to new data and the data are used to estimate values for the rate of flow of phloem sap through the insect and for the concentration required to prevent further excretion. Model predictions provide a close qualitative match to field data. However, rates of fluid flow from the trees through the insects would need to be much greater than the model assumes to produce values for droplet production that match field data. This suggests that the insects are playing an active role in regulating the flow rate. [Copyright &y& Elsevier]

In order to assess if ants attracted to honeydew-producing Hemiptera switched food sources when offered an additional, considerably rich and abundant food source, we studied the interaction between the plant Solanum lycocarpum (Solanaceae), an efn-lacking shrub, the predominantly diurnal ant Camponotus crassus (Hymenoptera: Formicidae), and two species of membracids (Enchenopa spp.; Homoptera: Membracidae). The study was done during September of 1999 at the Reserva Ecológica of IBGE (ca. 35 km S of Brasilia-DF, Brasil). The results show that ants were attracted both to the experimental food source and to the membracids. Moreover, the mean number of ants visiting the experimental nectary was significantly higher than those visiting the membracids. However, the membracids were never left unattended, which suggests that ants, even though presented with an abundant and rich food source, do not abandon other sources. The latter can be explained due to an increment in the honeydew production rate by the membracids, thus the attention levels of ants does not vary Another possibility is the fidelity of worker ants to a food source and its location. To show that ants switch food sources based on quality and/or quantity, and not merely based of their presence and/or absence, requires future research, both for plants with and without extrafloral nectaries. [ABSTRACT FROM AUTHOR]

The first samples of recently produced fir honeydew honey, obtained from Marchalina hellenica (Gennadius) (Homoptera, coccoidea, Margarodidae), originating from Mount Helmos in Greece were analysed and examined on the basis of physical and chemical parameters including humidity, acidity, sugar profile, hydroxymethylfurfural (HMF) and diastase activity. These data were compared with five other Mediterranean honeydew honeys of different origin obtained from the literature. Analytical data of the Greek fir honey from M. hellenica honeydew complied with the E.U. legal limits. The sensorial analysis indicated that the organoleptic characteristics of this type of honeydew honey were very similar to the Greek fir honey derived from other coccids with respect to colour, taste and smell. The statistical analysis of the physico-chemical parameters of the six honey types revealed two major groups. One group consisted of three Greek honeys and the other included one Spanish and two Italian honeys. it also showed a marked similarity between the two Greek honeys that came from M. hellenica honeydew on either fir or pine trees. [ABSTRACT FROM AUTHOR]

In studies of foraging behaviour in a multitrophic context, the fourth trophic level has generally been ignored. We used four aphid hyperparasitoid species:Dendrocerus carpenteri(Curtis) (Hymenoptera: Megaspilidae),Asaphes suspensusWalker (Hymenoptera: Pteromalidae),Alloxysta victrix(Westwood) (Hymenoptera: Alloxystidae) andSyrphophagus aphidivorus(Mayr) (Hymenoptera: Encyrtidae), to correlate their response to different cues with their ecological attributes such as host range and host stage. In addition, we compared our results with studies of primary parasitoids on the same plant–herbivore system. First, the olfactory response of females was tested in a Y-tube olfactometer (single choice: plant, aphid, honeydew, parasitised aphid, aphid mummy, or virgin female parasitoid; dual choice: clean plant, plant with aphids, or plant–host complex). Second, their foraging behaviour was described on plants with different stimuli (honeydew, aphids, parasitised aphids, and aphid mummies). The results indicated that olfactory cues are probably not essential cues for hyperparasitoid females. In foraging behaviour on the plant, all species prolonged their total visit time and search time as compared to the control treatment (clean plant). OnlyA. victrixdid not react to the honeydew. Oviposition in mummies prolonged the total visit time because of the long handling time, but the effect of this behaviour on search time could not be determined. No clear correlation between foraging behaviour and host stage or host range was found. In contrast to specialised primary aphid parasitoids that have strong fixed responses to specific kairomones and herbivore-induced synomones, more generalist aphid hyperparasitoids seem to depend less on volatile olfactory stimuli, but show similarities with primary parasitoids in their use of contact cues while searching on a plant. [ABSTRACT FROM AUTHOR]

Host feeding is the consumption of host tissue by the adult female parasitoid. We studied the function of destructive host feeding and its advantage over non-destructive feeding on host-derived honeydew in the whitefly parasitoid Encarsia formosa Gahan (Hymenoptera: Aphelinidae). We allowed parasitoids to oviposit until they attempted to host feed. We either prevented or allowed host feeding. Parasitoids had access to sucrose solution, with or without additional access to honeydew. Parasitoids that were allowed to host feed did not have a higher egg load 20 or 48 h after host feeding than parasitoids prevented from host feeding. Host feeding did not increase the number of eggs matured within these periods, nor did the time spent host feeding positively affect any of these response variables. On the other hand, the presence of honeydew did have a positive effect on egg load 20 and 48 h after host feeding compared with parasitoids deprived of honeydew. Parasitoids with access to honeydew matured more eggs within these periods than honeydew-deprived parasitoids. Host feeding increased life expectancy, but this effect was nullified when honeydew was supplied after the host-feeding attempt. In conclusion, feeding on honeydew could be an advantageous alternative to host feeding in terms of egg quantity and longevity. This applies especially to parasitoids exploiting Homoptera, because these parasitoids can obtain honeydew from the host itself. It is possible that destructive host feeding has evolved to enable females to sustain the production of high-quality anhydropic eggs, which may be important in the parasitoid's natural environment. We argue that future studies should take natural alternative food sources into more consideration. [ABSTRACT FROM AUTHOR]

This report documents the ongoing invasion of the Tokelau atolls by the long-legged ant, Anoplolepis gracilipes (Jerdon). These ants were collected from two of the three Tokelau atolls. On the island of Fenua Fala of Fakaofo Atoll, long-legged ants appear to be a recent arrival and occur in only a small area around one of the two ports. Most of the inhabited islands of Vao and Motuhuga on Nukunonu Atoll have been invaded, in addition to several of the uninhabited, forested islands. Despite this ant having been previously recorded from at least one island of Fakaofo and Nukunonu, these appear to be new invasions. Densities of up to 3,603 A. gracilipes per pitfall trap were caught per 24 hr. A significant reduction in ant species diversity was observed with increasing A. gracilipes densities. Densities of this ant were not uniformly high, perhaps due to variation in food availability. Prey such as crabs, ant colonies, and other insects were directly observed being attacked, and long-legged ants were observed to feed on honeydew produced by high densities of aphids, mealybugs, and scale insects on a variety of plants. Interspecific competition was investigated as an additional mechanism for the successful invasion. Long-legged ants found and removed bait faster than the dominant resident ant species, Paratrechina longicornis (Latreille), in forested areas of Nukunonu Island, though needing smaller numbers of recruits to achieve this result. This A. gracilipes invasion is of serious concern for the biodiversity of Tokelau and probably many of the other Pacific islands where these ants have invaded. [ABSTRACT FROM AUTHOR]