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The invasive mealybug Delottococcus aberiae De Lotto (Hemiptera: Pseudococcidae) has become a key pest in Mediterranean citrus. This mealybug species excretes honeydew that can be consumed by ants, which may give rise to mutualistic relationships and increase the invasibility of this exotic pest. Here, we studied the interaction between D. aberiae and ants in 16 citrus orchards located in the main citrus-growing area of Spain (Valencia) during two consecutive years. Four native ant species were observed feeding on D. aberiae honeydew, and Lasius grandis Forel (Hymenoptera: Formicidae) represented more than 95% of these ants. Lasius grandis was observed attending D. aberiae throughout the year and in all the orchard sites where the mealybug was present. Mealybug colony size had a positive effect on both relative and absolute ant-attendance by L. grandis, but these interactions varied across seasons. Moreover, we found a positive correlation between L. grandis activity and D. aberiae density in all citrus orchards. Our results suggest that the native ant L. grandis and the invasive pest D. aberiae have established a mutualistic association that promotes the establishment and accelerate the invasion of D. aberiae in Spanish citrus orchards. Further research with the aim of evaluating whether the management of this ant species can improve the control of D. aberiae in citrus would be recommended. [ABSTRACT FROM AUTHOR]

Simple Summary: Like many aphids, the sorghum aphid (SA) Melanaphis sorghi produces honeydew, a waste product that can increase parasitoid retention and elicit foraging behaviors on plants. We determined the potential of SA honeydew to retain a common aphid parasitoid, Aphelinus nigritus, to assess the practicality of SA biological control on grain sorghum, Sorghum bicolor. Since SAs also feed on Johnson grass, Sorghum halepense, an alternative plant host, we characterized the composition of honeydew from aphids feeding on either grain sorghum or Johnson grass and evaluated A. nigritus preference for honeydew produced on either plant species. Our study found that A. nigritus often remained on honeydew produced by SAs feeding on both grain sorghum and Johnson grass. However, despite sharing similar sugar, amino acid, and organic acid profiles, honeydew produced on Johnson grass was preferred by A. nigritus over honeydew produced on grain sorghum. Our results suggest that SA honeydew could facilitate A. nigritus parasitoid retention on Johnson grass to lower SA populations before the grain sorghum growing season. How aphid parasitoids of recent invasive species interact with their hosts can affect the feasibility of biological control. In this study, we focus on a recent invasive pest of US grain sorghum, Sorghum bicolor, the sorghum aphid (SA), Melanaphis sorghi. Understanding this pest's ecology in the grain sorghum agroecosystem is critical to develop effective control strategies. As parasitoids often use aphid honeydew as a sugar resource, and honeydew is known to mediate parasitoid–aphid interactions, we investigated the ability of SA honeydew to retain the parasitoid Aphelinus nigritus. Since SAs in the US have multiple plant hosts, and host–plant diet can modulate parasitoid retention (a major component in host foraging), we measured SA honeydew sugar, organic acid, and amino acid profiles, then assessed via retention time A. nigritus preference for honeydew produced on grain sorghum or Johnson grass, Sorghum halepense. Compared to a water control, A. nigritus spent more time on SA honeydew produced on either host plant. Despite similar honeydew profiles from both plant species, A. nigritus preferred honeydew produced on Johnson grass. Our results suggest the potential for SA honeydew to facilitate augmentation strategies aimed at maintaining A. nigritus on Johnson grass to suppress SAs before grain sorghum is planted. [ABSTRACT FROM AUTHOR]

Nylanderia fulva (Mayr) (Hymenoptera: Formicidae), or tawny crazy ant, is an invasive ant from South America that is spreading in the southern US. Extremely large populations of this ant can inundate urban and natural landscapes, and efficient control methods are lacking. This study was conducted to determine if activity of N. fulva would decline after controlling the hemipteran honeydew-producers they were tending. Potted plants infested with cottony cushion scale (Icerya purchasi Maskell) (Hemiptera: Monophlebidae) were treated with 2 imidacloprid insecticide formulations, and changes in N. fulva survival and foraging behavior were monitored. There were fewer ant trails, lower trailing intensity, less foraging, and less nesting in potted plants treated with either product than in the control pots. The use of low application rates of systemic insecticide to reduce honeydewproducing hemipterans, such as cottony cushion scale, could be an important component of the integrated pest management of N. fulva. [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]

In order for successful biological pest control using parasitoids, the most suitable naturally available food resources, as well as the fitness of parasitoids feeding on these resources, need to be ascertained. The goal of this study was to improve the understanding of the ecological requirements and consequent fitness of one such parasitoid, Telenomus podisi Ashmead ( Hymenoptera: Platygastridae), a predominant stink bug ( Heteroptera: Pentatomidae) egg parasitoid native to southeastern USA. We assessed the effects of carbohydrate resources readily available in the wasp's natural habitat on longevity and fecundity of female T. podisi under laboratory conditions. Carbohydrate diets included in the study were buckwheat, Fagopyrum esculentum Moench ( Polygonaceae) nectar and honeydew from cowpea aphid, Aphis craccivora Koch ( Hemiptera: Aphididae), compared with honey or water controls, which are normally used for laboratory rearing of this species. Eggs of Podisus maculiventris Say ( Hemiptera: Pentatomidae) were provided as hosts for oviposition. Honeydew of cowpea aphid proved to be as good a nutrition source as buckwheat nectar, in enhancing wasp longevity and fecundity. We also assessed the importance of honeydew freshness on the biological attributes of T. podisi, as honeydew becomes crystallized or highly viscous on drying up and therefore may pose an issue for uptake by parasitoids. We found that fresh honeydew of the pea aphid, Acyrthosiphon pisum Harris ( Hemiptera: Aphididae) significantly enhanced wasp longevity when compared to a diet of 1-day-old honeydew and the fava bean leaf (control). However, fresh honeydew of green peach aphid, Myzus persicae Sulzer ( Hemiptera: Aphididae) had the same effect on wasp longevity, fecundity, and proportion of male progeny as radish leaf (control). The potential of certain ubiquitously available food resources, in enhancing biological control of economically damaging pests by parasitoids, is being highlighted in this study; information which could be valuable in similar parasitoid-host systems as well. [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]

Conservation biological control involving the polyphagous aphid parasitoid, Aphidius colemani Viereck, may include provisioning resources from a variety of plant sources. The fitness of adult A. colemani was enhanced with the provision of food resources such as floral nectar from a range of both native and introduced plant species and aphid honeydew under laboratory conditions. However, enhanced fitness appeared to be species specific rather than associated with the whether the plant was a native or an introduced species. Parasitoid survival and fecundity were enhanced significantly in response to the availability of floral nectar and honeydew compared to the response to available extrafloral nectar. These positive effects on the parasitoid’s reproductive activity can improve the effectiveness of conservation biological control in nursery production systems because of the abundance and diversity of floral resources within typical production areas. Additionally, surrounding areas of invasive weeds and native vegetation could serve as both floral resources and honeydew food resources for A. colemani. [ABSTRACT FROM AUTHOR]

Aphid honeydew is one of the main energy sources for various ants in the temperate zone, nevertheless relatively little is known about the organization of the work of honeydew foragers (aphid milkers). This study focuses on the honeydew collecting strategies used by different ants in steppe and forest multi-species communities in Western Siberia. The behaviour of marked foragers of 12 species (Formica - 7, Lasius - 2, Camponotus - 1, Myrmica - 2) was recorded. Depending on the degree of the aphid milker specialization and degree of protection of the aphids five honeydew collecting strategies of various complexity were distinguished: unspecialized foragers in (I) "unprotected" aphid colonies (attended by ants <60% of time) and (II) "protected" colonies (attended >95% of time); (III) low "professional" specialization (ants "on duty" constantly attending aphid colonies); (IV) medium and (V) high "professional" specialization (clear division of tasks: honeydew collecting by "shepherds" and protection of trophobionts by "guards"; and honeydew transportation by "transporters" in V). Task specialization of the honeydew foragers is facultative: different ant taxa demonstrate a certain range of the honeydew collecting strategies of different complexities (Formica - I-V, Lasius - I-II, Camponotus - III, Myrmica - I-II) depending on the needs of their colony. The strategy used by ants did not depend on the species of aphid attended, but is strongly dependent on the species of ant, their colony size, available food resources and seasonality. In summer, the aphid milker specialization becomes more complex as ant colony size increases at both intra- and inter-specific levels and when food is scarce. In autumn Formica s. str. ants, which have the most advanced foraging strategy, adopt a simpler honeydew collecting strategy. Overall, the variety of honeydew foraging strategies seems to reflect the unequal contribution of different ants in forming trophobiotic interactions with aphids. [ABSTRACT FROM AUTHOR]

The introduction of invasive social wasp species of the genus Vespula ( Hymenoptera: Vespidae) to New Zealand has caused a major ecological problem, particularly in the beech forests ( Nothofagus spp.) of the South Island, where they have destabilized the native bird and invertebrate biodiversity. New attractants are under investigation as part of a search for pest management solutions. Fermenting brown sugar has been previously reported as a social wasp attractant. This work was undertaken to identify compounds from fermented brown sugar attractive to social wasps. Raw fermented brown sugar was confirmed to be attractive in a field trial and 10 chemical compounds present in the headspace were positively identified by coupled gas chromatography-mass spectrometry and synthetic references. During electroantennogram experiments, 3-methyl-1-butanol, 3-methylbutyl acetate, and ethyl hexanoate elicited high electrophysiological responses from Vespula vulgaris ( L.) antennae. These compounds mediated attraction of V. vulgaris wasps in forest margins by trapping. A blend of these compounds could be used as a lure in a monitoring tool, or even a local suppression method if combined with a toxin. [ABSTRACT FROM AUTHOR]

Mutualistic interactions between ants and Hemiptera are mediated to a large extent by the amount and quality of sugar-rich honeydew produced. Throughout the neotropics, the predaceous fire ant Solenopsis geminata ( Fabricius) ( Hymenoptera: Formicidae) is found in association with colonies of the pineapple mealybug, Dysmicoccus brevipes ( Cockerell) ( Hemiptera: Pseudococcidae), which they actively tend and protect from attack by natural enemies. In this study, we evaluate the effects of access to a sucrose solution on the mutualistic association between S. geminata and D. brevipes. Ten colonies of either species were established, with D. brevipes maintained on pumpkin, Cucurbita maxima Duchesne ( Cucurbitaceae), in screen cages. Five of the S. geminata colonies were permitted access to vials with 20% sucrose solution and a pumpkin with 20 adult mealybugs. The remaining ant colonies were allowed access to mealybug-infested pumpkins. Ant colonies with access to the sucrose solution attended mealybugs significantly less than those without additional sugar sources. Mealybug survival rates were similar under both treatments. Total body sugars and fructose were nearly twice as high in ants with access to honeydew and sucrose vs. those with access to honeydew and water. Fructose accumulated on the pumpkins over time in both treatments, suggesting that honeydew was not fully exploited by the ants. In conclusion, D. brevipes enjoy lower degrees of ant attendance when S. geminata have alternative sources of carbohydrates. We further discuss the significance of these findings for the conservation of predaceous ants and mealybug biological control. [ABSTRACT FROM AUTHOR]

Nylanderia fulva (Mayr) (Formicidae) is an invasive pest ant that has been reported in Florida, Texas, Louisiana and Mississippi. Workers tend various honeydew producing hemipterans in Florida landscapes and natural areas. We sought to understand the seasonal foraging activities of N. fulva and its relationship with honeydew producing hemipterans. Twenty-two hemipteran species were collected from 15 different plant hosts sampled in Florida from Jul 2010 to Jul 2012. The relative density of hemipterans on 4 plant species [live oak (Quercus virginiana Mill.; Fagales: Fagaceae), holly (Ilex cornuta Lindl.; Aquifoliales: Aquifoliaceae), magnolia (Magnolia grandiflora L.; Magnoliales: Magnoliaceae), and sugarberry (Celtis laevigata Willd.; Urticales: Ulmaceae)] was compared to the relative density of N. fulva at the base or stem of the selected plants. The number of N. fulva and hemipterans on each plant species was positively correlated from spring through fall. Both N. fulva and hemipteran populations increased from May to Sep and decreased from Oct to Apr. In addition, Cinara juniperivora (Wilson) (Hemiptera: Aphididae) on southern red cedar and Toumeyella liriodendri (Gmelin) (Hemiptera: Coccidae) on magnolia were covered with carton shelters presumably constructed by N. fulva, suggesting that this ant potentially protects certain hemipterans species. [ABSTRACT FROM AUTHOR]

The arboricolous dolichoderine ant Liometopum microcephalum (Panzer, 1798) is considered to be mainly predatory, although there are some reports of it tending aphids. The main objective of the present study was to confirm that this ant has a trophobiotic relationship with aphids and assess seasonal differences in its utilization of honeydew. We hypothesized that the worker ants on trees where they have their nest (nest tree) and trees where they are foraging (foraging trees) should differ in gaster mass and sugar content depending on their direction of movement, and that both should be highest in spring. From spring to summer 2009, ascending and descending workers were collected from nest and foraging trees at a locality in South Moravia, Czech Republic. Mass of their gasters and their content of total and reducing sugars were measured using chemical (photometric) methods. Differences in gaster mass confirmed the flow of liquid food from foraging to nest trees, but there were no significant between-month differences. Contents of total and reducing sugars were positively correlated with gaster mass. The gasters of workers descending from foraging trees contained significantly more reducing sugars than those of workers descending or ascending nest trees. The content of reducing sugars was lowest at the beginning of the ants' activity period in April and highest in June, with a non-significant drop in July. Results for total sugars were similar, with the decrease in July being significant. The concentration of sugars in the gasters of workers ascending and descending nest trees did not differ significantly but the absolute content of total sugars was higher in the gasters of ascending ants. Results from foraging trees confirmed that the ants collected the honeydew from these trees. Possible reasons for the ambigous results for nest trees are discussed. We conclude that trophobiosis is an important component of the nutritional biology of L. microcephalum. [ABSTRACT FROM AUTHOR]

Parasitoid survival and fecundity is generally enhanced with access to carbohydrate food sources. In many agricultural ecosystems, there is often a scarcity of suitable carbohydrates for parasitoids. This study compared the suitability of aphid honeydew and buckwheat nectar as diet for the aphid parasitoid Lysiphlebus testaceipes. Wasp lifespan and egg load were both increased with access to carbohydrates, but no differences were detected between the various carbohydrates diets tested. As aphid honeydew is a sufficient source of nutrition and L. testaceipes is a short-lived species, adding additional sources of carbohydrates like floral nectar strips to the agricultural landscape is unlikely to significantly increase the biological control exerted by L. testaceipes. [ABSTRACT FROM AUTHOR]

Abstract: Protopulvinaria pyriformis (Cockerell) (Hemiptera: Coccidae), a scale native to South America, is a major pest of laurel, Laurus nobilis L., and a potential pest of other ornamental plants in urban green areas. We evaluated the potential of Metaphycus helvolus (Compere) (Hymenoptera: Encyrtidae), its major parasitoid, as a biological control agent against P. pyriformis. This parasitoid was imported from South Africa to control soft scales in Europe more than 20years ago. Three factors limited its efficacy: (i) the scale frequently thwart parasitoid attack by removing its stylet and wiggling, a unique defense in soft scales, (ii) scale honeydew extended M. helvolus lifespan little more than water, and (iii) P. pyriformis did not encapsulate Metaphycus spp. eggs but immature parasitoids died during their prepupal stage, which may have been caused by laurel plant secondary compounds from laurel. This parasitoid mortality had not been previously reported in immature parasitoids of genus Metaphycus. These three novel aspects should be considered when choosing a suitable parasitoid in future soft scale biological control programs. [Copyright &y& Elsevier]

Honeydew-excreting hemipterans, such as mealybug pests, can be protected from their natural enemies by tending ants in return for honeydew, thereby compromising the aims of biological control. In this respect, antagonistic interactions between the ant Tapinoma nigerrimum, native to the Mediterranean basin, and the main natural enemies of both the vine mealybug (VMB), Planococcus ficus, and the citrus mealybug (CM), Planococcus citri, were assessed in laboratory conditions. Parasitism of vine and CMs by their respective parasitoids, Anagyrus sp. nr. pseudococci and Leptomastix dactylopii, was negatively affected by the ant T. nigerrimum. Similarly, T. nigerrimum was shown to significantly disrupt the predatory potential of ladybird larvae, Cryptolaemus montrouzieri, when foraging on host CMs. By contrast, the presence of the ant did not negatively influence the predatory activity of C. montrouzieri adults when feeding on CMs. Consequently, the encyrtid parasitoids A. sp. nr. pseudococci and L. dactylopii and the larval stage of the predator C. montrouzieri may be considered as T. nigerrimum-sensitive, whereas the adults of C. montrouzieri may be regarded as T. nigerrimum-resistant predators. Accordingly, the ant T. nigerrimum constitutes a threat to the biological control of mealybugs by either the encyrtids A. sp. nr. pseudococci and L. dactylopii or the larval stage of the ladybird C. montrouzieri. Hence, adequate control of T. nigerrimum is highly recommended before any release of these mealybugs' natural enemies. [ABSTRACT FROM PUBLISHER]

Honeydew from phloem-feeding insects and fruit fly insecticidal baits may serve as adult food resources for some insect species. In California (USA) olive orchards, the black scale [ Saissetia oleae (Olivier) (Hemiptera: Coccidae)] is a common honeydew producer, and spinosad-based fruit fly bait (GF-120) is used to control the olive fruit fly [ Bactrocera oleae (Rossi) (Diptera: Tephritidae)]. We investigated the effects of black scale honeydew and GF-120, as food resources, on adult foraging behaviour and survival of the olive fruit fly and two parasitoids in the olive agroecosystem: Scutellista caerulea (Fonscolombe) (Hymenoptera: Pteromalidae), a natural enemy of black scale, and Psyttalia humilis (Silvestri) (Hymenoptera: Braconidae), a parasitoid of the fruit fly. In food choice tests, female flies did not show a preference between GF-120 bait and honeydew, whereas male flies and the parasitoids we tested preferred honeydew. Adults of the three insect species readily fed on honeydew, but the natural enemies never ( P. humilis) or rarely ( S. caerulea) fed on GF-120 bait. Olfactometer tests further confirmed that the tested natural enemies were not attracted to GF-120 bait. The presence of honeydew significantly reduced fruit fly mortality when both honeydew and GF-120 were provided, compared with GF-120 given alone. A single meal of honeydew increased longevity in all insect species tested. The mean longevities of honeydew-fed insects were not significantly different from those feeding on clover honey. Our results suggest that the presence of honeydew would benefit the three insect species and may reduce the efficacy of GF-120 for fruit fly control because of preference of honeydew rather than fruit fly bait as a food resource. [ABSTRACT FROM AUTHOR]

Invasions by non-native insects can have important ecological impacts, particularly on island ecosystems. However, the factors that promote the success of invaders relative to co-occurring non-invasive species remain unresolved. For invasive ants, access to carbohydrate resources via interactions with both extrafloral nectary-bearing plants and honeydew-excreting insects may accelerate the invasion process. A first step towards testing this hypothesis is to determine whether invasive ants respond to variation in the availability of carbohydrate resources, and whether this response differs from that of co-occurring, non-invasive ants. We investigated the effect of carbohydrate subsidies on the short-term foraging and hemipteran-tending behaviours of the invasive ant Anoplolepis gracilipes (Formicidae) and co-occurring ant species on an extrafloral nectary-bearing plant by experimentally manipulating carbohydrate levels and tracking ant recruitment. We conducted experiments in 2 years at two sites: one site was invaded by A. gracilipes prior to 2007 and the other became invaded during the course of our study, allowing pre- (2007) and post-invasion (2009) comparisons. Short-term increases in carbohydrate availability increased the density of A. gracilipes workers on plants by as much as 400% and reduced tending of honeydew-excreting insects by this species by up to 89%, with similar responses across years. In contrast, ants at the uninvaded site in 2007 showed a weak and non-significant forager recruitment response. Across all sites, A. gracilipes workers were the only ants that responded to carbohydrate manipulations in 2009. Furthermore, ant-carbohydrate dynamics at a site newly invaded by A. gracilipes quickly diverged from dynamics at uninvaded sites and converged on those of the site with an established invasion. These findings suggest that carbohydrate resources may be particularly important for A. gracilipes invasions, and underscore the importance of species interactions, particularly putative mutualisms, in facilitating exotic species invasions. [ABSTRACT FROM AUTHOR]

As food items can vary in nutrient composition, diet selectivity can be an important trait to optimize nutrition. Most parasitoids depend as adults on various sugar-rich food sources such as nectar and honeydew. Nectar is often superior as a food source to honeydew, yielding higher parasitoid longevity. Little is known, however, of a preference parasitoids may have for one of these sugar sources and how this affects variation in their life-history traits. In this study, a combined empirical and modelling approach was adopted. First, it was investigated whether parasitoids show a feeding preference for nectar or honeydew in the laboratory and whether feeding history affects the preference of parasitoids. Second, using a process-based spatially explicit simulation model, it was evaluated how preference for nectar affects parasitoid life-history variables in fields with varying flower densities. The laboratory study revealed that naïve parasitoids showed no preference for either nectar or honeydew. However, when parasitoids had fed previously on honeydew and were then given the choice between nectar and honeydew, they preferred feeding on nectar to honeydew (with marginal statistical significance). Model simulations indicated that when nectar sources were present in the field, parasitoids that showed preference for nectar had a 1.3–1.8 times longer life span and parasitised up to 1.4 times more hosts than parasitoids that did not distinguish between honeydew and nectar. These combined empirical and modelling data suggest that aphid parasitoids may be able to discriminate between nectar and honeydew, and that a preference for nectar may enhance their longevity and reproduction in fields where nectar sources are available. [ABSTRACT FROM AUTHOR]

Extrafloral nectaries (EFNs) and hemipteran honeydew are liquid food rewards (FRs) that encourage ant visitation in many plant families in a wide variety of habitats. In this study we explored the diversity, distribution and interactions of exudate-gathering ants at three different liquid food rewards: nectar from EFNs on Croton sarcopetalus and honeydew from the aphids Aphis spiraecola and Dysaphis foeniculus on Eupatorium hookerianum (Asteraceae) and Foeniculum vulgare (Apiaceae) respectively. For each FR we measured ant diversity and performed quantitative and qualitative comparisons among ants associated with the FRs. In addition, a linear regression was performed to test for possible associations between aphid and ant abundances in the case of honeydew FRs. Eight out of the 23 ant species found fed on both nectar from EFNs and honeydew from aphids, four of which fed at all FRs. Two ant species visited both aphid species and 13 were found exclusively at either one of the FRs. Brachymyrmex brevicornis was the most abundant ant species and Pheidole sp.2 had the greatest occurrence. Both ant species diversity and richness were higher at EFNs of C. sarcopetalus. Regressions showed positive significant association between ants and aphids abundances both on E. hookerianum and F. vulgare. We can conclude that the three liquid food rewards compared here showed modest similarity in their ant fauna. Furthermore, there was selectiveness of ants towards EFNs of C. sarcopetalus, which might be due to food source attributes rather than co-evolutionary factors. [ABSTRACT FROM AUTHOR]

Abstract: Honeydew produced by aphids is a well-studied food source for mutualistic ants. Although considerable amounts drop onto the soil surface there are hardly any studies dealing with the potential effects of this additional C source for soil organisms, particularly those living at the soil surface. We studied this question by simulating honeydew and investigating plots under Populus canescens formerly infested with aphids on two ruderal sites differing in soil organic matter content and vegetation cover. The simulated honeydew input increased microbial biomass with some delay in week 10. Both natural and simulated honeydew raised the activity densities of an epigeic Collembola taxon, the Bourletiellidae, but not of the dominant Hemisotoma thermophila. The honeydew effect varied over time and with the site, and was more pronounced at the nutrient-poor site for Collembola, but not for microbial biomass. Ant consumption reduced the amount of honeydew reaching the soil surface by 50%. The activity density of H. thermophila was negatively related to ants and spiders, hinting at a top-down control. Honeydew clearly acts as a bottom-up force for soil organisms, which is however only part of the complex network connecting the food webs of primary producers and decomposers. [Copyright &y& Elsevier]

Feeding on sugar-rich foods such as nectar and honeydew is important for survival of many adult parasitoids. Especially in agricultural systems, honeydew is often the most prevalent carbohydrate source. However, relative to plant nectar, honeydew may be relatively unsuitable, as a result of an unfavourable sugar composition or the presence of secondary plant compounds. We studied survival of the aphid parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae) on honeydew collected from various aphid species feeding on potato ( Solanum tuberosum L., cv. Desiree) (Solanaceae), wheat ( Triticum aestivum L., cv. Bobwhite) (Poaceae), or artificial diet, as well as the sugar composition of the different honeydews. Honeydews from the tested aphid species on potato, wheat, or artificial diet were found to be relatively suitable food sources for adult A. ervi, although not always as suitable as a 2 M sucrose solution. There were differences in honeydew sugar composition among the different aphid species on the various host plants. Multivariate statistics showed that the factor ‘aphid species’ had a significant influence on the sugar composition of the honeydew, explaining 27% of the variation in the potato system and 89% in the wheat system. When exploring the relationship between carbohydrate composition of the honeydews from aphids on potato and wheat plants, and their nutritional value for A. ervi, data revealed that differences in parasitoid longevity can to some extent be explained by carbohydrate composition. Furthermore, our results confirm that sucrose and its hexose components glucose and fructose are very suitable carbohydrate sources for hymenopteran parasitoids and show that parasitoid survival on an equimolar solution of the two monosaccharides glucose and fructose does not exceed performance on the disaccharide sucrose. [ABSTRACT FROM AUTHOR]

Abstract: The effect of five different food resources (Coccus hesperidum honeydew, honey-water, Lobularia maritima, Fagopyrum esculentum and water only) on the longevity, fecundity, and progeny fitness of female Gonatocerus ashmeadi, an egg parasitoid of the glassy-winged sharpshooter (Homalodisca vitripennis) was investigated in the laboratory. Female G. ashmeadi survived 208–405% longer (up to 11.7 days longer) when fed C. hesperidum honeydew, honey-water, L. maritima and F. esculentum compared with water only. Similarly, all food treatments resulted in up to 378% more progeny (up to 80 more total progeny) than those females provisioned with water only. Mean percentage female progeny ranged from 70 to 81% and did not significantly differ between food treatments. The highest daily fecundity (22–29 progeny) occurred on the first day of oviposition for each food treatment and generally declined as female G. ashmeadi aged. Females deposited 61–100% of offspring within the first 5 days, after which, daily fecundity usually declined to 0–6 offspring per day. For all experimental diets except L. maritima, the number of male progeny oviposited remained relatively constant over time (∼2 males per oviposition bout) but the proportion of males oviposited increased as female G. ashmeadi aged, as oviposition rates declined and fewer female eggs were subsequently laid. There was a strong positive correlation between hind tibia length (HTL) and total progeny produced by G. ashmeadi fed honey-water. There was no correlation between HTL and longevity of female G. ashmeadi fed honey-water. Mean HTL of G. ashmeadi progeny did not significantly differ between food treatments. Estimated progeny fecundity based on HTL, indicated that offspring produced by females provided with water only was 12–15 eggs or 9–12% lower than females that had access to food resources. This difference was not statistically significant. Female progeny that were oviposited by females 9–13 days of age with access to food resources were significantly larger and contained on average 14–17 more eggs than those female progeny resulting from eggs deposited by females 1–8 days of age. [Copyright &y& Elsevier]

Abstract: The effect of dietary supplements on the longevity of male and female Gonatocerus ashmeadi, G. triguttatus and G. fasciatus (Hymenoptera: Mymaridae), was determined in the laboratory. Treatments included: water only, 3:1 honey–water solution, floral and extra-floral nectars from five different plants (excised stems from Fagopyrum esculentum, Lobularia maritima, Phacelia tanacetifolia, Anethum graveolens and Vicia faba), honeydew from Coccus hesperidum and Homalodisca vitripennis (formally H. coagulata), a commercially available food supplement (Eliminade) and citrus foliage. Additionally, the sugar composition of each food resource was determined using HPLC and whole flower extracts. Honey–water and F. esculentum nectar significantly increased longevity of male and female G. ashmeadi, G. triguttatus, and G. fasciatus up to 1860%, 1323% and 1459%, respectively, when compared with water. For both sexes and all three parasitoid species, survival on citrus foliage, H. vitripennis excrement, and P. tanacetifolia flowers was equivalent to that on water only. The longevity of G. ashmeadi and G. triguttatus was up to 539% higher on Eliminade compared with water only, however there was no significant effect of Eliminade on survival of G. fasciatus. Coccus hesperidum honeydew increased survival times up to 665% for all mymarid species compared with citrus foliage alone. HPLC analysis indicated that food resources most beneficial to Gonatocerus parasitoids possessed a high proportion of glucose (up to 44%) and fructose (up to 53%), suggesting that sucrose may not be as important for parasitoid survival. Citrus and P. tanacetifolia flowers contained favorable proportions of glucose and fructose, but the inability of Gonatocerus spp. to benefit from this may be related to flower morphology which could prevent access to nectar. [Copyright &y& Elsevier]

1. Many arthropods depend on carbohydrate-rich food sources such as nectar or honeydew. Nevertheless, we often know little about the extent to which various sugar sources contribute to the diet of arthropods. 2. One way to study food use in the field is to analyse guts of collected insects for source-specific compounds. Sugar sources often show distinct differences in their carbohydrate composition. This applies especially to honeydew, the excretion product of phloem-feeding ‘Sternorrhynchae’, which often features a broad range of phloem-feeder synthesized di- and oligosaccharides. 3. Out of these oligosaccharides, melezitose, has been widely used as an indicator of honeydew consumption. The use of melezitose or other honeydew saccharides as ‘signature sugars’ hinges on the assumption that the production of these sugars is unique to honeydew-secreting insects. 4. Here we show that the hymenopteran parasitoids Diadegma semiclausum and D. insulare synthesize the trisaccharides melezitose and erlose as well as the disaccharide maltose when fed sucrose, but not when feeding on an equimolar glucose–fructose mixture. 5. The presence of melezitose, erlose and maltose was confined to the parasitoid's digestive tract, indicating that enzyme activity is restricted to this area. D. semiclausum excrement contained low overall sugar concentrations and low relative levels of melezitose, erlose and maltose. 6. Possible functions of sugar synthesis in these nectar-feeding insects are discussed. [ABSTRACT FROM AUTHOR]

In response to herbivory by insects, several plant species have been shown to produce volatiles that attract the natural enemies of those herbivores. Using a Y-tube olfactometer, we investigated responses of the aphid parasitoid Diaeretiella rapae MacIntosh (Hymenoptera: Aphidiidae) to volatiles from Arabidopsis thaliana Columbia (Brassicaceae) plants that were either undamaged, infested by the peach-potato aphid, Myzus persicae Sulzer (Homoptera: Aphididae), or mechanically damaged, as well as to volatiles from just the aphid or its honeydew. In dual-choice experiments, female D. rapae given oviposition experience on A. thaliana infested with M. persicae were significantly attracted to volatiles from A. thaliana infested with M. persicae over volatiles from undamaged A. thaliana and similarly were significantly attracted to plants that had been previously infested by M. persicae, but from which the aphids were removed, over undamaged plants. Diaeretiella rapae did not respond to volatiles from M. persicae alone, their honeydew, or plants mechanically damaged with either a pin or scissors. We conclude that an interaction between the plant and the aphid induces A. thaliana to produce volatiles, which D. rapae can learn and respond to. Poor responses of D. rapae to volatiles from an A. thaliana plant that had two leaves infested with M. persicae, with the two infested leaves being removed before testing, suggested the possibility that, at this stage of infestation, the majority of volatile production induced by M. persicae may be localized to the infested tissues of the plant. We conclude that this tritrophic interaction is a suitable model system for future investigations of the biochemical pathways involved in the production of aphid-induced volatiles attractive to natural enemies. [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]