The impact of spruce aphids on nutrient flows in the canopy of Norway spruce

1 This study investigated the effects of honeydew from aphids in the canopy of Norway spruce (Picea abies (L.) Karst.) on the nitrogen chemistry of throughfall using a rainfall simulation experiment. Throughfall collected beneath infested trees was compared with that from beneath uninfested trees, while standardizing the quality and quantity of the precipitation and plant age. 2 Honeydew excreted by Cinara pilicornis (Hartig) and C. costata (Zett.) significantly increased the concentrations of dissolved organic carbon (DOC) and hexose-C in throughfall. The average concentrations of nitrogenous compounds (NH₄-N, NO₃-N) in throughfall collected beneath infested trees were significantly lower than beneath uninfested trees. 3 Multiple regression analysis indicated that the amount of rain and NH₄-N concentrations were the best predictors of the concentrations of dissolved organic nitrogen (DON) in throughfall. Parameters that were closely associated with the level of infestation (DOC, hexose-C concentrations) did not have a direct relationship with DON. About 40% of the reduction in the concentration of DON in the throughfall was attributed to aphid–micro-organism interactions. 4 Particle amino nitrogen (PAN)-concentrations were highest under infested trees in July after aphid numbers had declined, indicating a concomitant decline in microbial biomass after honeydew becomes a limiting resource. 5 The comparison of the concentrations of different nitrogen compounds in throughfall of infested and uninfested trees indicated that aphids affect the carbon and nitrogen cycles in the phyllosphere by providing energy that fuels the metabolism of the micro-organisms. These processes seem to occur very rapidly. 6 We discuss the significance of the results and the prospects of linking the ecology of micro-organisms and herbivores with flows of nutrients through the canopy of trees.     

Effects of Elatobium abietinum on nutrient fluxes in Sitka spruce canopies receiving elevated nitrogen and sulphur deposition

1 In a field acid mist simulation experiment, Sitka spruce (Picea sitchensis) was sprayed with different pollutant treatments: N, NH₄NO₃; S, Na₂SO₄; NS Acid, NH₄NO₃ + H₂SO₄ and control, no spray. Treatment effects on the abundance of the green spruce aphid Elatobium abietinum and honeydew production were assessed. In addition, needles were sampled for phyllosphere micro‐organisms. In a manipulative experiment, shoots were established and maintained as with or without E. abietinum infestation in order to determine the effects of infestation on needle loss and throughfall nutrient fluxes. 2 Aphid numbers were highest during the end of May and early June, with almost twice as many needles infested in the NS Acid treatment compared with the other treatments. Honeydew production was not affected by the treatments. 3 On infested shoots, increasing numbers of yellowing and dead needles were recorded above the throughfall collectors as the season progressed. The numbers of dead needles falling into the collectors were significantly higher beneath infested shoots. There were strong positive correlations between aphid numbers above the throughfall collectors, the number of yellowing and dead needles on the shoots and the number of needles in the funnels of the throughfall collectors. Litter production was more affected by aphid number than by pollutant treatment. 4 Bacteria, yeasts and filamentous fungi were more prolific on infested needles and treatment effects on colony forming units (CFUs) were most pronounced in the NS Acid treatment. 5 Fluxes of inorganic nitrogen beneath infested shoots were generally lower than beneath uninfested shoots. This effect was more pronounced in those treatments that supplied N i.e. N, NS Acid. The combination of aphid infestation and N‐addition exerted the strongest influence on nutrient fluxes. The fluxes of potassium and of organic carbon (DOC) were higher beneath infested shoots in all treatments, through most of the survey period.     

Linking aphid honeydew, throughfall, and forest floor solution chemistry of Norway spruce

In a seminatural manipulation experiment with artificial irrigation we followed throughfall and forest floor solution chemistry collected underneath aphid infested and uninfested Norway spruce. Solutions underneath infested trees showed significantly higher concentrations of dissolved organic carbon (DOC) but lower concentrations of dissolved organic nitrogen (DON), NO₃-N, and NH₄-N in throughfall solutions and of NH₄-N in forest floor solutions. Average concentrations were 40.5% (DON), 27.5% (NO₃-N), and 46.2% lower (NH₄-N) underneath infested trees in throughfall solutions, and 19.5% (DON), 9.4% (NO₃-N), and 42.0% (NH₄-N) lower in forest floor solutions. Differences in throughfall were more pronounced than in forest floor leachates. It is likely that honeydew is fuelling the metabolism of micro-organisms and thus critically affects above and below ground nutrient cycles. We emphasize the importance of linking the biology of herbivores and micro-organisms with geochemical processes.     

Insect herbivores and the nutrient flow from the canopy to the soil in coniferous and deciduous forests

Phytophagous insects can have severe impacts on forested ecosystems in outbreak situations but their contribution to flows of energy and matter is otherwise not so well known. Identifying the role of phytophagous insects in forested ecosystems is partly hindered by the difficulty of combining results from population and community ecology with those from ecosystem ecology. In our study we compared the effects of aphids and leaf-feeding lepidopterous larvae on the epiphytic micro-organisms in the canopies of spruce, beech and oak, and on the vertical flow of energy and nutrients from the canopies down to the forest floor. We particularly searched for patterns resulting from endemic herbivory rather than outbreak situations. Excreta of lepidopterous larvae and aphids promoted the growth of epiphytic micro-organisms (bacteria, yeasts, filamentous fungi) on needles and leaves, which suggests that micro-organisms were energy limited. Leachates from needles and leaves of infested trees contained higher concentrations of dissolved organic C and lower concentrations of NH₄-N and NO₃-N, relative to uninfested trees. The seasonal abundance of herbivores and micro-organisms significantly affected the dynamics of throughfall chemistry; for instance, concentrations of inorganic N were lower underneath infested than uninfested trees during June and July. There was little difference between the chemistry of soil solutions collected from the forest floor beneath infested and uninfested trees. Thus, under moderate to low levels of infestation the effects of above-ground herbivory seems to be obscured in the soil through buffering biological processes.     

Aphid honeydew and its effect on the phyllosphere microflora of Picea abies (L.) Karst

Aphids of the genus Cinara, feeding on Norway spruce, excrete copious amounts of honeydew, a carbon-rich waste product, which accumulates locally on needles and twigs. We investigated the role of honeydew as a potential source of energy which might promote the growth of micro-organisms in the phyllosphere of conifer trees. To approach this question, we followed the population dynamics of Cinara spp. in a natural forest stand over two seasons. We also studied the amounts of honeydew produced by individual aphids and identified potential parameters which might influence honeydew production. Finally, we determined the growth of micro-organisms on infested and uninfested needles of Norway spruce during the growing season. Confined to Picea abies, the investigated Cinara species only became abundant in midsummer, when needles and shoots were expanding. The populations showed only a single peak in abundance, the timing and magnitude of which may vary from year to year due to weather conditions, changes in plant quality in a yearly cycle or the impact of natural enemies. The amount of honeydew produced by individual aphids was dependent on the developmental stage of the aphid, the nutritional supply of its host plant and on the developmental state of the Norway spruce (e.g. bud burst, end of shoot extension). The presence of honeydew significantly increased the growth of bacteria, yeast and filamentous fungi on the surface of needles and there was a pronounced seasonal trend, with the highest abundance in midsummer correlating with the period of peak aphid abundance. Taken together, these findings indicate that aphids have an influence on microbial ecology in the phyllosphere of trees. The implication of our study, from interactions at the population level to effects and potential consequences for C and N fluxes at the level of forest ecosystems, is discussed.     

Females of Cotesia vestalis,a parasitoid of diamondback moth larvae,learn to recognise cues from aphid‐infested plants to exploit honeydew

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

Aphid-host plant interactions: does aphid honeydew exactly reflect the host plant amino acid composition?

Plants provide aphids with unbalanced and low concentrations of amino acids. Likely, intracellular symbionts improve the aphid nutrition by participating to the synthesis of essential amino acids. To compare the aphid amino acid uptakes from the host plant and the aphids amino acid excretion into the honeydew, host plant exudates (phloem + xylem) from infested and uninfested Vicia faba L. plants were compared to the honeydew produced by two aphid species (Acyrthosiphon pisum Harris and Megoura viciae Buckton) feeding on V. faba. Our results show that an aphid infestation modifies the amino acid composition of the infested broad bean plant since the global concentration of amino acids significantly increased in the host plant in response to aphid infestations. Specifically, the concentrations of the two amino acids glutamine and asparagine were strongly enhanced. The amino acid profiles from honeydews were similar for the two aphid species, but the concentrations found in the honeydews were generally lower than those measured in the exudates of infested plants (aphids uptakes). This work also highlights that aphids take large amounts of amino acids from the host plant, especially glutamine and asparagine, which are converted into glutamic and aspartic acids but also into other essential amino acids. The amino acid profiles differed between the host plant exudates and the aphid excretion product. Finally, this study highlights that the pea aphid, a “specialist” for the V. faba host plant, induced more important modifications into the host plant amino acid composition than the “generalist” aphid M. viciae.     

Fluxes and concentrations of dissolved organic carbon and nitrogen – a synthesis for temperate forests

Dissolved organic carbon (DOC) and nitrogen (DON)represent an important part of the C and N cycles inforest ecosystems. Little is known about the controlson fluxes and concentrations of these compounds insoils under field conditions. Here we compiledpublished data on concentrations and fluxes of DOC andDON from 42 case studies in forest ecosystems of thetemperate zone in order to evaluate controls on alarger temporal and spatial scale. The focus was onannual fluxes and concentrations in throughfall,forest floor leachates and soil solutions. In allcompartments considered, concentrations and fluxesdiffered widely between the sites. Highestconcentrations of DOC and DON were generally observedin forest floor leachates and in A horizons. Highestfluxes occurred in forest floor leachates. The fluxesof DOC and DON in forest floor leachates increasedwith increasing annual precipitation and were alsopositively related to DOC and DON fluxes withthroughfall. Variation in throughfall fluxes couldexplain 46% and 65% of the variation in DOC and DONfluxes from the forest floor, respectively. No generaldifference in DOC and DON concentrations and fluxes inforest floor leachates was found when comparingconiferous and hardwood sites. Concentrations of DOCin forest floor leachates were positively correlatedto the pH of the forest floor. Furthermore, there wasno relationship between organic C and N stocks, soilC/N, litterfall or mineral N inputs and concentrationsand fluxes of DOC and DON in forest floor leachates.Including all compartments, fluxes of DOC and DON werehighly correlated. Ratios of DOC to DON calculatedfrom fluxes from the forest floor were independent ofthe amount of annual precipitation, pointing to asimilar response of DOC and DON to precipitationconditions. A decrease in the ratio of DOC to DON withsoil depth as observed on a plot-scale, was notconfirmed by data analysis on a large scale. Thecontrols observed on annual fluxes and concentrationsof DON and DOC at regional scale differed from thosereported for smaller time and space scales.     

The effect of previously infested spruce needles on the growth of the green spruce aphid, Elatobium abietinum, and the effect of the aphid on the amino acid balance of the host plant

The effect of chlorosis induced in needles of Sitka and Norway spruce by the green spruce aphid on growth of the aphid is investigated, and the effect of infestation of the aphid on amino acid levels in Sitka spruce foliage is reported. On both Sitka and Norway spruce green spruce aphids were heavier when reared on chlorotic (previously infested) needles than when reared on green (previously uninfested) needles. The effect was more pronounced on Sitka than on Norway spruce. Infestation of the aphid altered the amino acid balance of Sitka spruce foliage but not the concentration of total amino acids. Possible causes of chlorosis, the influence of individual amino acids on aphid growth, the potential effect of chlorosis on outbreaks of the aphid and the differences in susceptibility of Sitka and Norway spruce to damage by the aphid are discussed.     

Performance of the green spruce aphid, Elatobium abietinum (Walker) on previously defoliated Sitka spruce

Abstract 1 The green spruce aphid, Elatobium abietinum, is an important defoliator of Sitka spruce in the U.K. However, it is usual for years in which high E. abietinum populations occur to be followed by a year with low aphid densities. The possibility that the performance of E. abietinum is reduced on previously infested Sitka spruce, and that this is the cause of year‐to‐year fluctuations in population density, was investigated by comparing population development and the growth rate of individual aphids on experimentally defoliated trees. 2 Separate experiments were performed to determine whether aphid performance was reduced either in the autumn immediately after defoliation in the spring, or was reduced in the spring of the next year. Different rates of initial defoliation on trees used to test aphid performance were created by artificially infesting the trees with aphids in the spring before the experiments, and varying the time of infestation. 3 Population development and the mean relative growth rate (MRGR) of individual aphids on previously defoliated and undefoliated Sitka spruce did not differ significantly in the spring of the next year. No differences were observed in the nutrient content of the 1‐year‐old needles of previously defoliated or undefoliated trees at this time. 4 In the autumn and winter immediately after spring defoliation, aphid MRGR was significantly higher on trees that had been heavily defoliated earlier in the season compared with trees that had been lightly defoliated. However, the difference in MRGR decreased over the winter period. Nitrogen, phosphorous and potassium concentrations were 9.4–12.2% higher, at the beginning of the autumn, in the current year needles of heavily defoliated trees than in the current year needles of lightly defoliated trees. 5 The experiments indicate that high populations of E. abietinum in the spring do not induce any defensive mechanisms in Sitka spruce that adversely affect subsequent generations of the aphid. By contrast, the results suggest that high spring densities of the aphid improve the nutritional quality of the current year's foliage for autumn generations.     

Tri-trophic level interactions affect host plant development and abundance of insect herbivores

Understanding the interactions among plants, hemipterans, and ants has provided numerous insights into a range of ecological and evolutionary processes. In these systems, however, studies concerning the isolated direct and indirect effects of aphid colonies on host plant and other herbivores remain rare at best. The aphid Uroleucon erigeronensis forms dense colonies on the apical shoots of the host plant Baccharis dracunculilfolia (Asteraceae). The honeydew produced by these aphids attracts several species of ants that might interfere with other herbivores. Four hypotheses were tested in this system: (1) ants tending aphids reduce the abundance of other herbivores; (2) the effects of ants and aphids upon herbivores differ between chewing and fluid-sucking herbivores; (3) aphids alone reduce the abundance of other herbivores; and (4), the aphid presence negatively affects B. dracunculifolia shoot growth. The hypotheses were evaluated with ant and aphid exclusion experiments, on isolated plant shoots, along six consecutive months. We adjusted linear mixed-effects models for longitudinal data (repeated measures), with nested spatial random effect. The results showed that: (1) herbivore abundance was lower on shoots with aphids than on shoots without aphids, and even lower on shoots with aphids and ants; (2) both chewing and fluid-sucking insects responded similarly to the treatment, and (3) aphid presence affected negatively B. dracunculifolia shoot growth. Thus, since aphids alone changed plant growth and the abundance of insect herbivores, we suggest that the ant–aphid association is important to the organization of the system B. dracunculifolia-herbivorous insects.     

Oviposition preferences of Maculinea alcon as influenced by aphid (Aphis gentianae) and fungal (Puccinia gentianae) infestation of larval host plants

Abstract 1. The influence of infestation of the larval host plant Gentiana cruciata on the egg‐laying preferences of the xerophilous ecotype of Alcon Blue butterfly (Maculinea alcon) was studied in a semi‐dry grassland area (Aggtelek Karst Region, Northern Hungary). 2. We examined whether oviposition patterns of females differed when G. cruciata stems were uninfested compared with when they were infested by an aphid (Aphis gentianae) or a rust (Puccinia gentianae) species. 3. Females laid more than 90% of their eggs on fertile, uninfested G. cruciata stems, although these stems comprised only ～ 50% of the total stems available. Stems infested by aphids were similar to uninfested ones in properties that had a strong correlation with egg numbers, and yet there were significantly fewer eggs on infested stems than on intact ones. 4. Females never laid eggs on parts of Gentiana stems infested by aphids, and the presence of Lasius paralienus ants, which have a mutualistic interaction with Aphis gentianae, did not increase the repulsive effect of aphids. Infection of Gentiana by Puccinia did not influence the egg‐laying behaviour of females, even though the flowers and buds of infested stems exhibited a delayed development. 5. Aphid infestation can influence butterfly oviposition patterns through both direct and indirect effects. The presence of aphids directly excluded oviposition, but our data also indicated the possibility of an indirect effect of aphid infestation. Stems that had no aphids at the last egg counting, but were infested prior to it, had significantly fewer eggs than those that were never infested.     

Aphid‐tending ants on introduced fennel: can resources derived from non‐native plants alter the trophic position of higher‐order consumers?

1. Although plant invasions often reduce insect abundance and diversity, non‐native plants that support phytophagous insects can subsidise higher trophic levels via elevated herbivore abundance. 2. Here ant–aphid interactions on non‐native fennel on Santa Cruz Island, California are examined. Fennel hosts abundant, honeydew‐producing fennel aphids. The patchiness of fennel and the relative lack of honeydew‐producing insects on other plants at our study sites suggest that assimilation of fennel‐derived honeydew would increase the abundance and decrease the trophic position of the omnivorous, aphid‐tending Argentine ant. 3. To assess the strength of the ant–aphid interaction, a comparison of ant abundance on and adjacent to fennel prior to and 3 weeks after experimental aphid removal was performed. Compared with control plants with aphids, ants declined in abundance on and around fennel plants following aphid removal. At the habitat scale, pitfall traps in fennel‐dominated habitats captured more ants than in fennel‐free scrub habitats. 4. To determine if assimilation of aphid‐produced honeydew reduces the ant's trophic position, variation in δ¹⁵N values among ants, plants and other arthropods was analysed. Unexpectedly, δ¹⁵N values for ants in fennel‐dominated habitats were higher than those of arthropod predators from the same sites and also higher than those of ants from fennel‐free habitats. 5. Our results illustrate how introduced plants that support phytophagous insects appear to transfer energy to higher trophic levels via elevated herbivore abundance. Although assimilation of fennel‐derived honeydew did not appear to reduce consumer trophic position, spatial variation in alternative food resources might obscure contributions from honeydew.     

Comparison of chlorophyll and carotenoid concentrations among Russian wheat aphid (Homoptera: Aphididae)-infested wheat isolines

Russian wheat aphid, Diuraphis noxia (Mordvilko), feeding injury on 'Betta' wheat isolines with the Dn1 and Dn2 genes was compared by assessing chlorophyll and carotenoid concentrations, and aphid fecundity. The resistant Betta isolines (i.e., Betta-Dn1 and Betta-Dn2) supported similar numbers of aphids, but had significantly fewer than the susceptible Betta wheat, indicating these lines are resistant to aphid feeding. Diuraphis noxia feeding resulted in different responses in total chlorophyll and carotenoid concentrations among the Betta wheat isolines. The infested Betta-Dn2 plants had higher levels of chlorophylls and carotenoids in comparison with uninfested plants. In contrast, infested Betta-Dn1 plants had the same level of chlorophyll and carotenoid in comparison with uninfested plants. Our data provide essential information on the effect of D. noxia feeding on chlorophyll and carotenoid concentrations for Betta wheat and its isolines with D. noxia-resistant Dn1 and Dn2 genes.     

Influence of aphid honeydew on the foraging behaviour of Hippodamia convergens larvae

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

The use of honeydew in foraging for aphids by larvae of the ladybird beetle, Coccinella septempunctata L. (Coleoptera: Coccinellidae)

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

Effects of Russian wheat aphid infestation on barley plant response to drought stress

The influence of Russian wheat aphid ( Diuraphis noxia Mordvilko) infestation on the response of barley ( Hordeum vulgare L. ev Hazen) plants to drought stress was investigated. Fourteen-day-old plants were infested with eight apterous adult aphids, which were removed 7 days later with systemic insecticide. Leaves previously infested with aphids had lower relative water content, reduced stomatal conductance, more negative water potential, lower levels of chlorophyll and higher levels of amino-N, proline and glycinebetaine than corresponding leaves from uninfested plants. When water was withheld for a period of 7 days after aphids were removed, the relative water content of previously infested plants dropped steadily from 0.89 to 0.60, while the relative water content of uninfested plants remained at about 0.94 for the first 4 days of the drought stress period followed by a steady drop to about 0.77 by the end of the drought stress period. Leaf water potentials dropped steadily during the drought stress period in both previously infested (-1.14 to -1.91 MPa) and unin-fested (-0.54 to -1.52 MPa) plants. Analysis of glycinebetaine and proline levels at the end of the drought stress period indicated that leaves of previously infested plants accumulated lower levels of these solutes than leaves from uninfested plants. Upon alleviation of drought stress, plants previously infested with aphids showed little increase in dry weight while younger leaves and tillers from uninfested plants showed large increases. It is concluded that Russian wheat aphids cause drought-stress symptoms in leaves of infested plants even in the presence of ample root moisture. The observations of low levels of glycinebetaine and proline present in leaves after water was withheld from roots and lack of leaf growth upon alleviation of drought stress in previously-infested plants, suggest that aphid infestation limits the capacity of barley plants to adjust successfully to drought stress.     

Effects of honeydew and insecticide residues on the distribution of foraging aphid parasitoids under glasshouse and field conditions

The opposing effects of attraction to host-derived kairomones and repellency from the pyrethroid insecticide deltamethrin were investigated with aphid parasitoids from the genus Aphidius (Hymenoptera: Aphidiinae). The spatial distribution of female parasitoids was recorded in a series of experiments conducted in a small glasshouse containing wheat plants either infested with cereal aphids, Sitobion avenae (F.) (Homoptera: Aphididae), uninfested or treated with the recommended field concentration of deltamethrin. The number of parasitoids per plant were counted at 0.5 h, 1 h and then at one hourly intervals up to 8 h after release. Parasitoids showed a strong aggregation response to aphid-infested plants compared to adjacent uninfested plants. With the introduction of insecticidetreated plants around the aphid-infested plants, parasitoids showed a greater tendency to disperse away, resulting in fewer parasitoids on plants and significantly lower rates of aphid parasitism. The degree of aphid fall-off from plants was a good indicator of parasitoid foraging activity. In field studies, using sticky traps to measure the activity of parasitoids in plots sprayed with water, deltamethrin and/or an artificial honeydew solution, repellent properties were evident for up to 2 days after application. The attraction/arrestment stimuli associated with the honeydew solution were sufficient for parasitoids to continue searching insecticide-treated areas. The implications of these findings for parasitoids searching crops contaminated with aphid-derived kairomones and insecticides are discussed.     

Pea aphids (Acyrthosiphon pisum Harris) reduce secretion of extrafloral nectar in broad bean (Vicia faba)

1. Herbivores sometimes suppress plant defences. This study tested whether the presence of pea aphids (Acyrthosiphon pisum Harris) on broad bean (Vicia faba) led to decreased secretion of extrafloral nectar (EFN) which functions as an indirect plant defence against herbivores. 2. To determine effects of aphid infestation on EFN secretion, a comparison was done between EFN secretion in uninfested plants and that in plants infested by A. pisum and another aphid species (Aphis craccivora Koch). 3. When broad bean plants were infested by A. pisum, they secreted significantly smaller amounts of EFN than did uninfested plants and A. craccivora‐infested plants. There was no significant difference in EFN secretion between uninfested plants and A. craccivora‐infested plants. The number of extrafloral nectaries did not differ among the three treatments. 4. These results suggest that A. pisum reduced EFN production in broad bean plants.     

Incidence of honeydew in southern pine-hardwood forests: implications for adult parasitoids of the Southern Pine Beetle,Dendroctonus frontalis (Coleoptera: Scolytidae)

The incidence of homopteran honeydew was measured under pines and hardwoods located within 10 Dendroctonus frontalis Zimmermann infestations on the Talladega National Forest (Alabama) during summer, 2000. There were more honeydew droplets beneath hardwood trees compared to pines, but no difference beween D. frontalis infested and uninfested pines. The number of honeydew droplets per tree differed between sampling dates, likely due to the effect of mean weekly temperature and rainfall on homopteran populations. The presence of honeydew on hardwoods may explain why D. frontalis parasitoids forage in areas that lack hosts and suggests that the fecundity of D. frontalis parasitoids may not be limited by adult nutrition.