Vertical stratification in the spatial distribution of the beech scale insect (Ultracoelostoma assimile) in Nothofagus tree canopies in New Zealand

Abstract. 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.     

Preliminary evidence suggests that beech scale insect honeydew has a negative effect on terrestrial litter decomposition rates in

Honeydew production by New Zealand beech scale insects (Ultracoelostoma spp., Hemiptera: Margarodidae) is widely considered to have a positive influence on native animal abundance and ecosystem functioning. As a first assessment of whether there is a positive relationship between honeydew throughfall and litter decomposition rates, we placed experimental litter bags in each of 10 high and 10 low honeydew plots in mixed southern beech (Nothofagus) forest at each of two sites, Lakehead and Rotoiti, in the Nelson Lakes National Park. High and low honeydew plots were selected using sooty mould biomass on the forest floor as a surrogate for honeydew throughfall, as sooty mould biomass was shown to be strongly correlated (r = 0.906) with scale insect population size at the Rotoiti site. Contrary to our expectation, terrestrial litter decomposition was significantly lower in high honeydew plots than in low honeydew plots, at both Lakehead and Rotoiti. The presence of introduced wasps (Vespula spp., Hymenoptera: Vespidae) at the Lakehead site did not appear to have any significant effect on litter decomposition rates, despite the fact that wasps are thought to intercept much of the honeydew produced in this forest. Variance in litter decomposition rates between high and low honeydew treatments was predominantly determined by a direct relationship between sooty mould biomass and litter decomposition rate at the scale of individual litter bags. However, the mechanistic explanation for the observed relationship is unclear. Future studies should be directed towards quantifying the functional relationship between honeydew throughfall and growth rates of sooty mould, and their subsequent effects on abiotic conditions, microarthropod community dynamics and microbial activity rates in litter.     

Separating host-tree and environmental determinants of honeydew production by Ultracoelostoma scale insects in a Nothofagus forest

Abstract 1. Sugar‐rich honeydew excreted (‘produced’) by insects feeding on phloem sap is a key energy flow in a range of temperate and tropical ecosystems. The present study measured honeydew produced by Ultracoelostoma sp. (Homoptera: Coelostomidiidae) scale insects feeding on Nothofagus solandri var. solandri (Hook f.) Oerst. trees in a temperate evergreen forest in New Zealand. Simultaneous measurements of environmental variables and canopy photosynthesis were conducted to allow separation of host‐tree and environmental determinants of honeydew production. These relationships were further examined in experiments where canopy photosynthesis was manipulated by shading or plant nitrogen levels increased by foliar spray. 2. Rates of honeydew production varied nine‐fold from a maximum (± 1 SE) of 64.4 ± 15.2 mg dry mass m⁻² bark h⁻¹ in early summer (December) to a minimum of 7.4 ± 4.2 mg m⁻² h⁻¹ in winter (August). Rates of production measured 1.4 m from the base of the trees’ stems varied significantly with stem diameter, and were higher on medium‐sized (18 cm diameter) than small or large stems. 3. Rates of production were significantly related to environmental conditions over the hours preceding measurement (air temperature and air saturation deficit averaged over the preceding 24 and 12 h respectively). There was no evidence that rates of production were directly related to short‐term changes in the supply of carbohydrates from the canopy (either when compared with measurements of unmanipulated photosynthetic rate, or after sugar levels were manipulated by shading 80% of host‐trees’ leaf area), or to changes in phloem nitrogen content. 4. The results show that there is no clear effect of host‐tree carbon supply on honeydew production; if production is related to photosynthesis, the effect of this is much less important that the large and significant direct effect of environmental conditions on honeydew production.     

Patterns in aphid honeydew production parallel diurnal shifts in phloem sap composition

Variation in phloem sap composition is important in determining aphid performance and is known to occur at both diurnal timescales and in response to plant age. For field grown potato plants, Solanum tuberosum L. (Solanaceae), we determined diurnal variation in components of phloem sap, measured by ethylene diamine tetra‐acetate exudation, and tested for impacts of plant age. The effects of plant age and diurnal cycles on honeydew production by Macrosiphum euphorbiae (Thomas) and Myzus persicae (Sulzer) (both Hemiptera: Aphididae) were also quantified. Both the ratio of sucrose to amino acids and the composition of amino acids in phloem sap varied significantly with time of day. Dietary essential amino acids contributed a smaller proportion of amino acids in the phloem sap of older plants and during early phases of the diurnal cycle. The only significant effect on aphid honeydew production was of the diurnal cycle for Ma. euphorbiae, although increased honeydew production during the day when compared with the production at night, was consistent across the two species. In contrast with studies carried out at seasonal scales, we found limited evidence for variation in phloem sap composition in response to plant age, consistent with our results for honeydew production. These data highlight the need for improved understanding of how seasonal and diurnal physiology of plants influence performance in phloem sap feeding insects.     

Feeding rates and carbohydrate metabolism by Bemisia tabaci (Homoptera: Aleyrodidae) on different quality phloem saps

Abstract.The effects of water stress on phloem sap quality of the melon, Cucumis melo, and how this, in turn, has an impact on the sweet potato whitefly, Bemisia tabaci were studied. Melon plants were grown under watering regimes that produced plants with or without water stress. Plants showed strong developmental responses to the treatments; water-stressed plants were shorter, with fewer, smaller leaves than those without stress. There was, however, no effect of plant water stress on the development period of whiteflies feeding on these plants, or on the weights of male or female adults. Honeydew production was used as an indirect measure to test whether the absence of insect developmental or behavioural effects was due to differential phloem sap ingestion. Feeding rates on the stressed plants were almost half those on unstressed plants, and there was also variation in the daily pattern of honeydew production. Phloem sap and honeydew were analysed to determine why the feeding behaviours differed. Amino acid composition of the phloem sap was similar in both groups of plants, but carbohydrate concentrations were greater in water-stressed plants, indicating that lower feeding rates may have been due either to the physical or nutritional quality of the phloem sap. The honeydew of insects that were feeding on water-stressed plants contained a greater concentration of carbohydrate than those on unstressed plants, and was composed of a significantly greater proportion of glucose and the disaccharide, trehalulose. This isomerization of more complex sugars from those in the diet suggests that B. tabaci uses a mechanism of osmoregulation to actively maintain its internal water status. It is concluded that transient conditions of water stress in this host plant do not affect the development of B. tabaci, due to physiological and behavioural changes in response to diets with different nutritional and physical properties. The implications of this finding for the feeding biology of B. tabaci on desert-grown crops are discussed.     

Effect of host-tree and environmental variables on honeydew production by scale insects (

Honeydew excreted by phloem-sap sucking scale insects (Ultracoelostoma sp.) living in the bark of beech (Nothofagus solandri:) trees growing at a high elevation (900 m) site in the Craigieburn range of Canterbury, New Zealand, was measured over four days during 1–10 May 1996. Average standing crop of honeydew sugar was 3.1 mg m-2, and ranged from 0.4 to 5.5 mg m-2. Daily production of honeydew sugar ranged from 0.2 to 1.5 mg insect-1 24 h-1, and 4.1 to 45.9 mg m-2 24 h-1. Honeydew production varied significantly between trees, and trees with the highest mean individual rates of production (mg sugar insect-1 24 h-1) tended to be those with the highest numbers of insects per unit bark area. Air temperature averaged over the 24 hours preceding each production sample explained 65% of the variability in 3-hourly honeydew production, showing that honeydew production is controlled by environmental and host-tree variables. Using this data, together with previously published estimates of carbon uptake and seasonal variability in honeydew standing crop, it was estimated that carbon contained in honeydew is equivalent to 1.8% of net primary production of beech trees at Craigieburn. Determining the effects of environmental variables on tree and insect physiology will allow formulation of an environmentally-driven process-based model of honeydew production.     

Phloem-sap feeding by animals: problems and solutions

The incidence of phloem sap feeding by animals appears paradoxical. Although phloem sap is nutrient-rich compared with many other plant products and generally lacking in toxins and feeding deterrents, it is consumed as the dominant or sole diet by a very restricted range of animals, exclusively insects of the order Hemiptera. These insects display two sets of adaptations. First, linked to the high ratio of non-essential:essential amino acids in phloem sap, these insects contain symbiotic micro-organisms which provide them with essential amino acids. For example, bacteria of the genus Buchnera contribute up to 90% of the essential amino acids required by the pea aphid Acyrthosiphon pisum feeding on Vicia faba. Second, the insect tolerance of the very high sugar content and osmotic pressure of phloem sap is promoted by their possession in the gut of sucrase-transglucosidase activity, which transforms excess ingested sugar into long-chain oligosaccharides voided via honeydew. Various other animals consume phloem sap by proxy, through feeding on the honeydew of phloem-feeding hemipterans. Honeydew is physiologically less extreme than phloem sap, with a higher essential:non-essential amino acid ratio and lower osmotic pressure. Even so, ant species strongly dependent on honeydew as food may benefit from nutrients derived from their symbiotic bacteria Blochmannia.     

Characterization and correlation of DC electrical penetration graph waveforms with feeding behavior of beet leafhopper, Circulifer tenellus

Feeding behavior of beet leafhopper, Circulifer tenellus (Baker) (Homoptera: Cicadellidae), was studied with a DC electrical penetration graph. Nine different electrical penetration graph waveforms associated with feeding were identified and characterized. Waveforms were correlated with specific feeding behaviors using a number of techniques, including high magnification video recording, honeydew analysis, stylectomy, and histological processing. Waveforms were grouped into three phases based on feeding behavior: pathway phase (waveforms A, B1, B2, and C), non-phloem ingestion phase (waveform G), and phloem phase (waveforms D1, D2, D3, and D4). No ingestion was found to occur during waveforms A, B1, B2, and C. Waveform G was associated primarily with ingestion of xylem sap and occasionally with ingestion of mesophyll sap. Stylet tips were located in phloem during waveforms D1, D2, and D3, and waveforms D2 and D3 were correlated with ingestion of phloem sap. Waveform D4 probably also plays a role in phloem ingestion, because D4 is very brief and always occurs embedded in either waveform D2 or D3. In contrast to most other homopteran insects, rate of honeydew production (and hence rate of ingestion) was much lower on phloem than on xylem. More rapid rates of ingestion are expected on phloem, because its high turgor pressure drives sap into the feeding insect whereas the negative pressure of xylem sap is expected to cause a slow rate of ingestion. The very slow ingestion rate of beet leafhopper feeding on phloem suggests that it is not able to exploit the high turgor pressure of phloem to achieve the high rate of ingestion that is typical of phloem ingestion by other insects.     

Soluble carbon production by honeydew scale insects in a New Zealand beech forest

We estimated the annual production of honeydew per unit land area of beech (Nothofagus spp.) forest by measuring the amount of honeydew produced in 24 h by scale insects (Ultracoelostoma spp.) (Hemiptera: Margarodidae) every month for 2 years. We used exclosures to prevent animals (notably Vespula wasps) removing honeydew, and we compared the standing crop of honeydew inside permanently closed exclosures with that outside exclosures. Honeydew production and the number of honeydew droplets was highly variable between individual trees, tree type, position on tree, and, exclosure type, and within and between years. The amount of honeydew available outside exclosures was significantly reduced in year 2, predominantly by Vespula wasps, even though wasp density was relatively low. Sugar composition also varied between tree type and between years. Up to 5% of the sugar was glucose, with varying proportions of fructose, sucrose and oligosaccharides. The surface area of trees infested with scale insects was estimated using allometric regression relationships between tree diameter and total surface area of tree trunk and branch material. These estimates were combined with measurements of tree diameter in 10-m radius circular plots to give a production estimate of between 3500 and 4500 kg dry weight honeydew ha^-1 year^-1. Using this data, combined with previously published estimates of carbon uptake, it was estimated that between 6 and 8% of net primary productivity was released as honeydew. Honeydew scale insects provide large amounts of biologically available carbon in the form of soluble sugar. It is a crucial resource for the above-ground system, and probably also for the below-ground system. We conclude that scale insects have the potential to function as keystone species in these forests.     

Competition for honeydew between two social wasps in South Island beech forests,New Zealand

Summary Honeydew is a sugary secretion of beech scale insects (Ultracoelostoma spp.). Two introduced species of wasps forage on the sugar droplets in New Zealand beech forests. We hypothesize that competition between them may explain whyVespula germanica has become locally extinct in beech (Nothofagus) forest invaded byV. vulgaris. Changes in behaviour of the two wasp species in response to changes in the honeydew resource were monitored. Foraging and rainfall separately and together reduced the honeydew standing crop. In response to the standing crop decreasing, more wasps were found on honeydew trees, they became less active, spent more time lapping the tree surface, and ingested drops of honeydew at a slower rate.V. vulgaris was more active, and obtained drops and energy at a higher rate thanV. germanica. These behavioural differences may lead to competitive advantages affecting queen size and possibly survival.     

Sap‐feeding insects on forest trees along latitudinal gradients in northern Europe: a climate‐driven patterns

Knowledge of the latitudinal patterns in biotic interactions, and especially in herbivory, is crucial for understanding the mechanisms that govern ecosystem functioning and for predicting their responses to climate change. We used sap‐feeding insects as a model group to test the hypotheses that the strength of plant–herbivore interactions in boreal forests decreases with latitude and that this latitudinal pattern is driven primarily by midsummer temperatures. We used a replicated sampling design and quantitatively collected and identified all sap‐feeding insects from four species of forest trees along five latitudinal gradients (750–1300 km in length, ten sites in each gradient) in northern Europe (59 to 70°N and 10 to 60°E) during 2008–2011. Similar decreases in diversity of sap‐feeding insects with latitude were observed in all gradients during all study years. The sap‐feeder load (i.e. insect biomass per unit of foliar biomass) decreased with latitude in typical summers, but increased in an exceptionally hot summer and was independent of latitude during a warm summer. Analysis of combined data from all sites and years revealed dome‐shaped relationships between the loads of sap‐feeders and midsummer temperatures, peaking at 17 °C in Picea abies, at 19.5 °C in Pinus sylvestris and Betula pubescens and at 22 °C in B. pendula. From these relationships, we predict that the losses of forest trees to sap‐feeders will increase by 0–45% of the current level in southern boreal forests and by 65–210% in subarctic forests with a 1 °C increase in summer temperatures. The observed relationships between temperatures and the loads of sap‐feeders differ between the coniferous and deciduous tree species. We conclude that climate warming will not only increase plant losses to sap‐feeding insects, especially in subarctic forests, but can also alter plant‐plant interactions, thereby affecting both the productivity and the structure of future forest ecosystems.     

Invasive wasps,not birds,dominate in a temperate honeydew system

The Avian Convergence Hypothesis states that avian–honeydew associations are likely to develop when biogeographic and/or climatic factors limit the formation of ant–honeydew associations (the dominant association in tropical ecosystems). In this study we examine a honeydew‐influenced forest system in an island archipelago where ant diversity is low but invasive Vespula wasp species (Vespidae) are present. We found honeydew production was highly seasonal, with both standing crop and 24‐h production peaking in summer. When Vespula wasps were abundant (summer and autumn) they preferentially visited infested trees and fed regularly on honeydew droplets on infested branches. Two ant species occasionally fed on honeydew. No other insects or birds were observed feeding on honeydew during the study period. With the exception of Vespula, honeydew does not appear to be a preferred food source in this community, possibly because of the range of other food resources available in surrounding forest, farmland and gardens. The abundance of Vespula wasps at the site may also have disrupted bird–honeydew associations. We suggest the Avian Convergence Hypothesis could be restated to explicitly include both nectar availability and invasive social insects as both are likely to influence bird use of honeydew.     

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

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

Determining the impact of scale insect honeydew, and invasive wasps and rodents, on the decomposer subsystem in a New Zealand beech forest

Relatively few studies have considered how aboveground invasive consumers influence decomposer communities. We investigated the potential effects of three types of animals on the decomposer subsystem in a floristically simple New Zealand Nothofagus forest. These animals are the native beech honeydew scale insect (Ultracoelostoma spp.) that secretes large amounts of sugar-rich honeydew that washes to the soil, invasive social wasps (Vespula spp.) that remove honeydew and prevent it from reaching the ground, and invasive rodents (the house mouse (Mus musculus) and ship rat (Rattus rattus)) that are predators of litter invertebrates. We performed a 4 years manipulative experiment involving addition of synthetic honeydew to the soil surface at amounts equal to that washed to the soil both in the absence and presence of wasps. All treatments were subjected to both exclusion and non-exclusion of rodents. Full honeydew addition influenced several components of the belowground community (both positively and negatively), and promoted fungi and fungal feeding fauna at the expense of bacteria and bacterial-feeders. The reduced addition of honeydew (representing effects of wasps) reversed some (but not all) effects of full honeydew addition. Rodents also influenced some belowground organisms, often reversing the effects of honeydew addition. The honeydew levels simulating wasp effects and the presence of rodents both greatly promoted humus carbon and nutrient storage relative to all other treatments, highlighting that invaders can alter soil carbon sequestration and nutrient capital. Our study points to invasive animals modifying the effects of a native animal on multiple components of the decomposer subsystem.     

Antifeedant effects of plant lectins and an enzyme on the adult stage of the rice brown planthopper,Nilaparvata lugens

The feeding activity of adult rice brown planthopper,Nilaparvata lugens (Stal) on selected proteins incorporated into artificial diets was examined, based on quantitative analysis of honeydew excretion, in order to determine the mechanism of action of antimetabolic proteins towards homopteran pests. The lectinsGalanthus nivalis agglutinin (GNA), wheat germ agglutinin (WGA) and the enzyme soybean lipoxygenase (LPO), previously demonstrated to be toxic to this insect, reduced honeydew excretion levels of adultNilaparvata lugens over a 24 h period when incorporated into artificial diet at 0.1% {w:v}, indicating that these proteins acted as antifeedants. Of the proteins tested GNA was the most effective antifeedant, reducing honeydew droplet production by 96%, although after 24 h there was some recovery in the honeydew excretion levels and thus the insects appeared to tolerate the presence of the antifeedant with time. The lectinPisum sativum agglutinin (PSA), previously demonstrated to be non-toxic, showed no antifeedant properties.     

Field evidence for indirect interactions between foliar‐feeding insect and root‐feeding nematode communities on Nicotiana tabacum

Abstract 1. As herbivory often elicits systemic changes in plant traits, indirect interactions via induced plant responses may be a pervasive feature structuring herbivore communities. Although the importance of this phenomenon has been emphasised for herbivorous insects, it is unknown if and how induced responses contribute to the organisation of other major phytoparasitic taxa. 2. Survey and experimental field studies were used to investigate the role of plants in linking the dynamics of foliar‐feeding insects and root‐feeding nematodes on tobacco, Nicotiana tabacum. 3. Plant‐mediated interactions between insects and nematodes could largely be differentiated by insect feeding guild, with positive insect–nematode interactions predominating with leaf‐chewing insects (caterpillars) and negative interactions occurring with sap‐feeding insects (aphids). For example, insect defoliation was positively correlated with the abundance of root‐feeding nematodes, but aphids and nematodes were negatively correlated. Experimental field manipulations of foliar insect and nematode root herbivory also tended to support this outcome. 4. Overall, these results suggest that plants indirectly link the dynamics of divergent consumer taxa in spatially distinct ecosystems. This lends support to the growing perception that plants play a critical role in propagating indirect effects among a diverse assemblage of consumers.     

The effect of β-aminobutyric acid on the growth of herbivorous insects feeding on Brassicaceae

dl ‐β‐Aminobutyric acid (BABA) is a nonprotein amino acid that can enhance defences in a variety of plants against a wide range of pathogens. BABA can also reduce infestation by phytopathogenic nematodes and has recently been shown to suppress the growth of aphids feeding on legumes. This investigation examined the effect of applying BABA as a root drench to a range of Brassicaceae, including Arabidopsis thaliana, on the performance of two species of aphid (Myzus persicae and Brevicoryne brassicae) and the larvae of two species of Lepidoptera (Trichoplusia ni and Plutella xylostella). Application of BABA reduced the performance of all four insect species, and inhibition of insects occurred on all the plants tested. The results illustrate that BABA‐induced resistance (BABA‐IR) can affect generalist and specialist insect herbivores and inhibit insects feeding with mandibulate as well as sap‐feeding mouthparts. The BABA‐induced suppression of B. brassicae and P. xylostella feeding on A. thaliana provides a means to further examine the mechanisms of BABA‐IR to insects using this model plant.     

Feeding responses of an oligophagous bean aphid, Megoura crassicauda, to primary and secondary substances in Vicia angustifolia

The feeding behaviour of the aphid Megoura crassicauda Mordivilko (Homoptera: Aphididae), which feeds selectively on plants in the genus Vicia (Fabaceae), was studied. The aphids deposited proteinaceous stylet sheaths intercellularly towards the phloem tissues of host plants. Similar stylet sheaths were formed on a Parafilm membrane when host‐specific acylated flavonoid glycosides [two 2″‐O‐(E)‐p‐coumaroyl esters of quercetin 3‐O‐diglycosides] present in the extracts of the narrow vetch, Vicia angustifolia L., were supplied in the solution covered by the membrane. In contrast, their corresponding deacyl analogues, present more abundantly in the host plant tissues, were not stimulatory, which suggested specificity in the structural requirements of the probing stimulants. While the aphids imbibed an artificial diet composed of primary nutrients (e.g., sucrose and amino acids) and produced a large quantity of honeydew, acylated flavonoids alone and non‐acylated flavonoids supplied with the nutrients more or less suppressed honeydew production. These findings implied that the acylated flavonoids serve as a cue to navigate the stylet sheath towards the phloem prior to sap‐sucking, whereas non‐acylated flavonoids may serve as a negative stimulus to refrain from sucking during tissue penetration before tapping the phloem, although the distribution of these compounds in the plant tissues remains unknown. Thus, the feeding behaviour of M. crassicauda appears to be controlled by multiple chemical stimuli in the process of the settling on its host plant.     

Distribution and abundance of endemic coelostomidiid scale insects (Hemiptera: Coelostomidiidae) in Auckland forests, New Zealand

Sap-feeding insects can excrete considerable quantities of sugar-rich honeydew. In New Zealand, South Island beech (Nothofagus spp.) forests are shaped by the extensive honeydew resource produced by two endemic coelostomidiid species (Ultracoelostoma assimile and U.?brittini) and geckos on northern islands are known to feed on the honeydew of a third endemic coelostomidiid, Coelostomidia zealandica. There are six other endemic coelostomidiid species in New Zealand that utilise a range of plant hosts but the ecological role of these species is poorly understood. A survey of mainland forests in the Auckland Ecological Region was conducted in February?April 2006 to investigate the distribution and abundance of coelostomidiids in this area. Three coelostomidiid species were detected in the survey (C.?zealandica, C.?pilosa and C.?wairoensis) and five new host?scale insect associations were identified. C.?zealandica was uncommon, C.?pilosa was widespread in broadleaved?podocarp forest but only formed light infestations, and C.?wairoensis was present in all teatree stands examined, often forming heavy infestations on k?nuka (Kunzea ericoides). Infested k?nuka trees had sooty moulds growing on them and exotic wasps were regularly seen feeding on C.?wairoensis honeydew. The extent and intensity of C. wairoensis infestation on k?nuka suggests it will have community-level impacts.     

Honeydew composition and its effect on life-history parameters of hyperparasitoids

1. Diets that maximise life span often differ from diets that maximise reproduction. Animals have therefore evolved advanced foraging strategies to acquire optimal nutrition and maximise their fitness. The free-living adult females of parasitoid wasps (Hymenoptera) need to balance their search for hosts to reproduce and for carbohydrate resources to feed. 2. Honeydew, excreted by phloem-feeding insects, presents a widely available carbohydrate source in nature that can benefit natural enemies of honeydew-producing insects. However, the effects of variation in honeydew on organisms in the fourth trophic level, such as hyperparasitoids, are not yet understood. 3. This study examined how five different honeydew types influence longevity and fecundity of four hyperparasitoid taxa. Asaphes spp. (Pteromalidae) and Dendrocerus spp. (Megaspilidae) are secondary parasitoids of aphid parasitoids and are thus associated with honeydew-producing insects. Gelis agilis and Acrolyta nens (both Ichneumonidae) are secondary parasitoids of species that do not use honeydew-producing hosts. 4. Most honeydew types had a positive or neutral effect on life span and fecundity of hyperparasitoids compared with controls without honeydew, although negative effects were also found for both aphid hyperparasitoids. Honeydew produced by aphids feeding on sweet pepper plants was most beneficial for all hyperparasitoid taxa, which can partially be explained by the high amount of honeydew, but also by the composition of dietary sugars in these honeydew types. 5. The findings of this study underline the value of aphid honeydew as a carbohydrate resource for fourth-trophic-level organisms, not only those associated with honeydew-producing insects but also ‘interlopers’ without such a natural association.