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.     

Seasonal changes in the distribution of green spruce aphid Elatobium abietinum (Walker) (Homoptera: Aphididae) in the canopy of Sitka spruce

1 Seasonal changes in the distribution of green spruce aphid Elatobium abietinum (Walker) within the canopy of 20–25‐year‐old Sitka spruce are described based on data from two low‐altitude sites (310–420 m above sea‐level), two mid‐altitude sites (500–550 m a.s.l) and one high‐altitude site (610 m a.s.l). 2 Aphids were counted throughout the canopy on shoots representative of all needle age‐classes present at each whorl of branches. Counts were made during the middle week of each month from September to July for 4 years (1999–2003), and mean E. abietinum densities at each canopy position were calculated separately for each month and for the low‐, mid‐ and high‐altitude sites. 3 During September to November, the highest densities of E. abietinum occurred on 3–4‐year‐old needles on branches low in the canopy. Over the winter and spring, the centre of the aphid’s distribution shifted outward and upward, so that by June of the next year the highest aphid densities occurred on current and 1‐year‐old needles on branches near the top of the tree. 4 The aphid distribution was re‐set each year during July, at the time when the nutrient quality of the host was in decline and E. abietinum populations were decreasing. Aphid densities decreased less on 3–4‐year‐old needles than on current and 1‐year‐old needles, suggesting that older needles were a superior food resource at this time of year and in the autumn. However, other factors, such as higher temperatures in the upper canopy during the summer or differential mortality caused by natural enemies, could also have contributed to the change in distribution. 5 The outward and upward shift in the aphid distribution over the winter period provided no evidence that aphids at positions lower and deeper in the canopy were better insulated from freezing temperatures and had higher over‐winter survival rates. Mean air temperatures at the top and bottom of the canopy during the winter were also found to differ by only 0.1–0.2 °C. 6 The percentage of the total aphids per tree that occurred on current or 1‐year‐old needles varied widely between seasons and between sites. Consequently, sampling programmes designed to estimate total population numbers of E. abietinum have little option but to sample needles throughout the canopy, and at regular intervals during the period when the aphid is abundant.     

Frequency and intensity of drought stress alters the population size and dynamics of Elatobium abietinum on Sitka spruce

The green spruce aphid, Elatobium abietinum, is the most important defoliating pest of Sitka spruce, Picea sitchensis, in the UK. Populations are expected to increase in response to predicted climate change, therefore placing Sitka spruce under increased risk of widespread and severe defoliation. The effect of spring–summer drought stress on E. abietinum population dynamics and development over multiple years was assessed in a field experiment under five different drought treatments with differing intensities and frequencies. The impact on host tolerance, in terms of needle retention, was also investigated. No differences in the length of time taken to reach peak population size were observed. Despite this, E. abietinum populations were found to respond positively to low amplitude intermittent stress, with high densities maintained for longer after the peak. Spring peak densities of aphids did not differ among drought levels, although an autumn peak was observed consistently over 2 years on trees subjected to continuous severe drought. This suggested advancement in the onset of Sitka spruce dormancy. Aphid infestation significantly increased percentage needle loss under all drought treatments, although no differences were observed between drought levels. An interaction between aphid presence and drought treatment was, however, observed during a second year of drought where no aphids were present. The study herein presented has contributed to the understanding of E. abietinum population responses under a changing climate. The implications for damage to host Sitka spruce are of relevance to forest management strategies, as an increase in drought events are predicted in the UK.     

The effects of high temperatures on individuals and populations of the green spruce aphid Elatobium abietinum (Walker)

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Defoliation of Sitka spruce by the European spruce sawfly, Gilpinia hercyniae (Hartig): a retrospective analysis using the needle trace method

Abstract 1 During 1968–74, an outbreak of the European spruce sawfly Gilpinia hercyniae, defoliated many stands of Sitka spruce in commercial forests in mid‐Wales. The needle trace method was used to determine retrospectively the temporal pattern and intensity of defoliation in a stand of Sitka spruce in Hafren Forest that had been damaged severely at the time of the sawfly outbreak. 2 An initial calibration experiment, designed to test the reliability of the needle trace method when applied to Sitka spruce, indicated that artificially induced defoliation of up to 75% was detected by the technique with an accuracy of ±7%. Higher rates of defoliation were underestimated by up to 17%. 3 For the main sample of trees, the needle trace method demonstrated that retention of needle sets (average needle retention) was reduced by up to 33–38% over a 10‐year period coincident with the G. hercyniae outbreak and the years immediately afterward when the canopy was recovering. 4 Analysis of needle loss within separate needle cohorts and age‐classes revealed that 1‐year‐old needles were the most severely defoliated. The density of 1‐year‐old needles was reduced by 51–78% in 1970–73. 5 Defoliation at the time of the G. hercyniae outbreak was associated with reductions in annual height, radial, and volume increments of 24–49%, 30–59% and 32–56%, respectively. Radial and volume increments suffered their greatest reductions 1 year later than height increments. 6 The study demonstrates that the needle trace method can be applied successfully to Sitka spruce to quantify defoliation caused by an insect outbreak occurring many years previously, and that the technique can provide data on needle loss that is valuable for interpreting reductions in tree growth.     

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.     

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.     

Drought intensity and frequency have contrasting effects on development time and survival of the green spruce aphid

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Consistency of resistance to attack by the green spruce aphid (Elatobium abietinum Walker) in different ontogenetic stages of Sitka spruce

Abstract 1 The susceptibility of different genotypes of 29‐year‐old Sitka spruce to damage by the green spruce aphid, Elatobium abietinum, was investigated in a progeny trial where aphid damage on individual trees had previously been assessed twice in an earlier stage of ontogenetic development. The progeny trial comprised 14 open‐pollinated families originating from a clonal seed orchard that had been established using mature spruce trees selected for aphid resistance. 2 Previous investigations had demonstrated that resistance was inherited by the offspring, and that differences in resistance between progenies of the individual orchard clones were highly significant. 3 Susceptibility to aphid attack was recorded as the percentage loss of previous year's needles. Differences in susceptibility recorded between the juvenile trees were found to persist after the trees had developed into the closed‐canopy, sexually reproducing stage. Needle loss of the families was significantly less than that of the reference population of Sitka spruce. 4 Hybrids between Sitka spruce and white spruce were defoliated more heavily than pure Sitka spruce, and the difference increased with age. 5 Family heritability of resistance was estimated as 0.60 compared to 0.73 when the trees were assessed in the juvenile stage. The genetic correlation based on family means between damage in the juvenile and sexually reproducing stand was high (0.83), indicating a high consistency of resistance to the aphid over years and ontogenetic stages. 6 A skewed distribution of defoliation indicated that major genes are involved in the expression of resistance, and that the genetics behind resistance has a nonadditive component.     

Season and drought stress mediate growth and weight of the green spruce aphid on Sitka spruce

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Damage caused by the Green Spruce Aphid to Norway and Sitka spruce needles

Three-month-old needles of Sitka spruce were less susceptible to Elatobium abietinutn than 15-month-old needles. Symptoms appeared after longer aphid feeding times but only a proportion of damaged needles fell. After short feeding periods symptoms appeared in more Norway spruce needles than in Sitka spruce, whereas longer feeding periods resulted in more needles producing symptoms in the Sitka spruce. The symptoms took 4—6 days longer to appear in Norway spruce, and needle fall followed a longer feeding period than on Sitka spruce. Following 72 h feeding, needle fall occurred more quickly on Sitka spruce than on Norway spruce. The time taken for needle fall to occur was inversely related to the feeding time in Sitka spruce but such a response was not evident in Norway spruce. The results are discussed in relation to the differences exhibited in the probing behaviour of the aphid on the two spruces.     

Genetic diversity of the green spruce aphid (Elatobium abietinum Walker) in north-west Europe

Abstract 1 The Random Amplified Polymorphic DNA (RAPD) method was used to investigate genetic diversity of anholocyclic populations of the green spruce aphid, Elatobium abietinum Walker, in north‐west Europe. 2 The results showed that the aphid in this region was divided genetically into three major groups. Aphids from the British Isles and north‐west France comprised the first group, the second group consisted of aphids from Denmark and Iceland, and the third group consisted of aphids from Norway. 3 The results indicated a significant level of gene flow within and between sites and geographical regions, especially in the British Isles and north‐west France. Lateral migration of the aphid and/or sexual reproduction is likely to have facilitated the gene flow. 4 The implications of these findings on management of the green spruce aphid are discussed.     

Effects of temperature on overwintering populations of the green spruce aphid Elatobium abietinum

Recorded minimum temperatures of – 7 ^oC or lower noticeably reduced overwintering populations of the green spruce aphid, Elatobium abietinum, in north-east Scotland. It is suggested that, at these temperatures, ice formation in the needles of the host Sitka spruce, caused attached aphids to freeze. Aphid mortality also occurred when maximum temperatures did not rise above + 6 ^oC for prolonged periods, possibly as a result of starvation following an extended chill coma. A diagrammatic representation of the main factors affecting anholocyclic populations of E. abietinum during the winter is presented to emphasize the governing role played by temperature. The balance between mortality and recruitment determines the size of the population at the end of the winter, and this in turn determines the subsequent summer infestation. It should be possible, therefore, to predict aphid outbreaks either from winter temperatures or from the number of aphids present at the end of the winter. Temperature records obtained from integrating thermometers indicated that the inside of the lower crown tended to be the warmest part of the tree during the winter, resulting in greater aphid survival in the lower branches.     

Supercooling and the low-temperature survival of the green spruce aphid Elatobium abietinum

Laboratory investigations into the low-temperature tolerance of the green spruce aphid, Elatobium abietinum, revealed that the insect was killed by freezing. Aphids and host Sitka spruce needles showed similar seasonal changes in supercooling ability. A noticeable increase in this ability occurred between June and October. Aphids were more susceptible to low temperatures when attached to the plant. It is suggested that mortality resulted from ice which formed in the sap of the host needles and spread into the feeding aphids via their mouthparts. Neither the chlorotic banding of needles, caused by aphid feeding, nor needle length affected needle supercooling. Increased duration of exposure increased the probability of freezing of supercooled needles at low temperatures. A small percentage of first-instar nymphs supercooled to much lower temperatures than the remainder of the population. These were newly born nymphs whose high supercooling ability markedly decreased when they began to feed.     

Reproduction of an arboreal aphid pest,Elatobium abietinum,is altered under drought stress

Climate change in the UK is predicted to increase both winter temperatures and the frequency of summer drought events. Elatobium abietinum, the green spruce aphid, is the most important defoliating pest of Sitka spruce, Picea sitchensis, a conifer very widely used in British forest industry. This aphid is expected to respond strongly to altered climate, with changes to population densities leading to more frequent serious outbreaks and defoliation. The impact of simulated spring–summer drought on the reproductive performance of E. abietinum was investigated under laboratory conditions. Rates were assessed under five drought treatments of differing frequencies and intensities to characterize the direction of responses under different drought scenarios, and in time‐staggered trials to explore seasonal variation. Variation in the response of reproduction to water deficit was mediated by drought frequency and magnitude. Low‐amplitude, moderate intermittent stress improved reproductive rates, while severe stress, both continuous and high‐amplitude intermittent, had a detrimental impact when compared with observations made on well‐watered controls. Season was also found to modify the response, with improvements to plant nutritional quality under high‐amplitude stress reflected by improving reproduction. Despite this, no differences in rates were found during the autumn, suggesting no advancement in spruce dormancy under drought. Drought stress therefore has the potential to alter E. abietinum population densities, structure and phenology in Sitka spruce plantations, with implications for forest management, damage levels and natural control of the aphid under future altered climate.     

Effect of open-air fumigation with sulphur dioxide and ozone on phyllosphere and endophytic fungi of conifer needles

The phyllosphere microbial populations inhabiting the needles of three conifer species, Scots pine (Pinus sylvestris L.), Sitka spruce (Picea sitchensis L.) and Norway spruce (Picea abies (L.) Karst.), exposed to SO₂ and O₃, in an open-air fumigation experiment were analysed over a 3 year period using serial dilution after washing, direct plating and a fluorescein diacetate (FDA) enzyme assay. Total fungal populations ranged from 10² to 10⁵ colonyforming units (CPU) g^?1 fresh weight of needles. The dominant fungi isolated from needles varied with tree species and isolation technique; Aureobasidium pullulans (de Bary) Arnaud was most common on Scots pine and Norway spruce and white yeasts on Sitka spruce using the dilution plating method. However, direct plating of needle segments onto culture media indicated that Sclerophoma pythiophila (Corda) Hohnel was dominant on Scots pine and A. pullulans on Sitka and Norway spruce. Green needles of Sitka spruce were found to be endophytically colonized by Rhizosphaera kalkhoffii Bubak, but seldom by Lophodermium piceae (Fuckel) Hohn during extensive sampling in 1990. Statistical analyses revealed significant differences (P<0.05) between plots in the 3 year mean of the total fungal populations or the fungal biomass (FDA assay) on all three tree species. Differences between plots were also observed for a number of dominant component species. Data were also analysed for treatment effects. A significant effect of SO₂ treatment was observed on the total fungal populations on Sitka spruce (P<0.05) which were reduced markedly by the low-SO₂ treatment, while the O₃ treatment caused a significant increase in total fungal numbers on Scots pine (P<0.05). The FDA activity on needles of both Scots pine and Sitka spruce was noticeably higher in the 0₃-only treatment plot, but the overall O₃ effect was not significant. Treatment effects were also detected on the occurrence of component species. The serial dilution method revealed an SO₂ effect (P<0.05) of a reduction in the occurrence of pink yeasts on Sitka spruce and an O₃ effect (P<0.05) of an increase in the occurrence of S. pythiophila on Sitka spruce (P<0.01) but a decrease of Epicoccum nigrum Link and Cladosporium spp. on Scots pine. The direct-plating method revealed an SO₂ effect of an increase in S. pythiophila on Norway spruce (P<0.05). Ozone treatment caused a significant increase in the isolation of a black strain of A. pullulans on Norway spruce (P<0.05). Endophytic colonization of Sitka spruce needles by R. kalkhoffii was found to be increased on two occasions by O₃ exposure.     

Differences in the probing behaviour of Elatobium abietinum feeding on Sitka and Norway spruces

The probing behaviour of Elatobium abietinum (Walk.) on the needles of Norway spruce and Sitka spruce was directly related to the stomatal frequency accompanied by heavy deposition of saliva, and multiple branching and divergence of the salivary sheath, all of which were associated with the impenetrability of the endodermis. On Norway spruce these features were very greatly reduced where there was a higher rate of probing coupled with a higher rate of penetration into the phloem, which was not accompanied by early uptake of phloem sap when compared with aphids on Sitka spruce. These differences in probing behaviour are related to damage symptoms exhibited by the host trees.     

Genetic differentiation of the green spruce aphid (Elatobium abietinum Walker), a recent invader to Iceland

1 The RAPD method (Random Amplified Polymorphic DNA) was used to investigate genetic diversity of the green spruce aphid, Elatobium abietinum Walker, a pest introduced recently to Iceland. 2 This aphid in Iceland comprised two polymorphic populations, one in the east and the other in the west of the country. The genetic variation between sites within a population was continuous and appeared to be in good agreement with geographical distances. 3 In the eastern population the variation was greater between sites than within sites, whereas in the western population the pattern of variation appeared to be the opposite. This overall greater genetic variation in the eastern population could be due to its having been colonized earlier than the western one. 4 The study also demonstrated a close relationship between the green spruce aphid in Iceland and aphids from Denmark, which agrees with their assumed origin. The differences in introduction time, adaptation and competitiveness between the two Icelandic populations are discussed.     

The effects of SO2 and O3 on the foliar nutrition of Scots pine, Norway spruce and Sitka spruce in the Liphook open-air fumigation experiment

Foliar elements were analysed in Scots pine, Sitka spruce and Norway spruce over a 6 year period before and during continuous exposure to SO₂ and O₃ in an open-air fumigation experiment. Sulphur dioxide treatment elevated foliar sulphur concentration in all species, and there were increases in foliar nitrogen in the two spruce species but not in pine. The concentrations of cations were frequently increased by SO₂ treatment, but there was no correlation between the sulphur concentration of needles and their total cation charge. SO₂-related elevations of foliar magnesium were correlated with the concentration of this element in soil solution, but the mechanism by which other cations were enhanced remains unclear. The only consistent effects on nutrient ratios were for SO₂ treatments to increase sulphur/cation ratios.     

Effects of altitude on aphid-mediated processes in the canopy of Norway spruce

Abstract 1 The abundance of aphids and their honeydew are important in shaping the ecology of food web interactions and nutrient cycling in forests of Norway spruce. Here, the effects of the different environmental conditions at two study sites located at different altitudes (500 m, 765 m a.s.l.), in the Fichtelgebirge, north‐eastern Bavaria, Germany, on the abundance of Cinara pilicornis and their influence on epiphytic microorganisms on shoots of Picea abies were compared. Subsequent changes in throughfall fluxes were measured over a period of 12 weeks beneath infested and reference trees. In a laboratory experiment, the effects of ultraviolet (UV) radiation on microbial mortality and C and N concentrations in leachates were determined. 2 The warmer and drier conditions at the low altitude site favoured an early onset to aphid multiplication in spring compared with the high altitude site, where aphid numbers peaked 3 weeks later. 3 The presence of honeydew was associated with a significant increase in the total number of cultured epiphytic filamentous fungi, yeasts and bacteria in 12 of the 18 sample units, indicating better culturability or growth, whereas altitude had no significant effect on cultured cell numbers. By contrast to the reference trees, the high dissolved organic carbon (DOC) and hexose‐C fluxes beneath infested trees at the peak in aphid abundance, in June and July, resulted in a concomitant decrease in the fluxes of total inorganic nitrogen beneath infested trees (low altitude: ?19.7%; high altitude: ?52.3%). Fluxes of organic nitrogen were significantly higher beneath infested trees at the time of infestation. Similarly, potassium fluxes in throughfall increased 1.6–2.0‐fold in response to aphid infestation. 4 The exposure of infested and uninfested shoots of Norway spruce to UV‐A and UV‐B radiation only weakly affected epiphytic microbial mortality and did not affect the concentrations of the different nitrogen compound in leachates. However, bacteria, tended to be more active in the leachates collected from infested shoots, which resulted in the higher concentrations of aminosugar‐N. The aphids had a more pronounced effect on the concentrations of DOC in leachates, with average DOC concentrations being 4.2‐fold higher than in leachates from uninfested shoots. 5 It is suggested that, even at very low densities, aphids exert a strong influence via honeydew on the performance of microorganisms, and nutrient and energy flow, in spruce forests.