Reproduction and Control of the Invasive Polyphagous Shot Hole Borer, Euwallacea nr. fornicatus (Coleoptera: Curculionidae: Scolytinae), in Three Species of Hardwoods: Effective Sanitation Through Felling and Chipping.

The invasive ambrosia beetle polyphagous shot hole borer is one member of the cryptic species, Euwallacea nr. fornicatus (Eichhoff), and poses a great ecological and environmental threat to ornamental and native hardwood trees and agriculturally important tree crops in southern California, United States. We monitored the emergence of polyphagous shot hole borer adults from chipped and unchipped cut logs of infested boxelder, Acer negundo L. (Sapindales: Sapindaceae), California sycamore, Platanus racemosa Nutt. (Proteales: Platanaceae), coast live oak, Quercus agrifolia Née (Fagales: Fagaceae), and red willow, Salix laevigata Bebb (Malpighiales: Salicaceae), every week for 4 to 5 mo. No polyphagous shot hole borer adults emerged from chipped or unchipped coast live oak logs, suggesting this species is not a preferred reproductive host. However, following chipping, a small number of polyphagous shot hole borer adults emerged from boxelder (up to 7 wk) and from California sycamore and red willow (both up to 9 wk). A significantly greater number of polyphagous shot hole borer adults emerged from unchipped logs of boxelder (up to 14 wk) and California sycamore and red willow (both up to 5 mo). Chipping of boxelder, California sycamore, and red willow by using common commercially available chippers reduced polyphagous shot hole borer emergence by over 97%. Emerged polyphagous shot hole borer adults were strongly female-biased, regardless of host. Chipping treatments were highly effective, but for the complete elimination of polyphagous shot hole borer from woody material, other sanitation measures such as solarization in conjunction with chipping are recommended. Environmental and ecological impact of polyphagous shot hole borer is reviewed and discussed.

The Effects of Weather on the Flight of an Invasive Bark Beetle, Pityophthorus juglandis.

The walnut twig beetle, Pityophthorus juglandis Blackman (Coleoptera: Scolytidae), vectors the fungus Geosmithia morbida, which has been implicated in thousand cankers disease of walnut. Little is known about the flight behavior of the insect across seasons, or about the variability in its flight patterns with weekly fluctuations in weather. We sampled flying adults weekly over a 142-week period (from 29 August, 2011 to 2 June, 2014) with 12-unit black plastic multiple funnel traps baited with a male-produced aggregation pheromone in California, USA. Up to 5000 beetles were captured per trap per week, although catches in most weeks were less than 100 insects. Trap catches were regressed against terms for precipitation, solar radiation, vapor pressure, air temperature, relative humidity, wind speed, and trap catches in preceding weeks. The number of beetles captured in each of the preceding two weeks explained most variation in a current week's catch. This strong temporal autocorrelation was present in regression models developed for males, females, and both sexes pooled. These models were improved by including two environmental variables. Captures of P. juglandis increased with mean weekly air temperature and decreased with increasing mean minimum relative humidity. The percentage of variation in male, female, or total trap catch explained by the temporal variables and the two environmental variables in these multiple regression models ranged from 72% to 76%. While the flight of this invasive insect will likely be affected by site-specific factors as it spreads to new areas, the strong temporal correlation present in this system may provide a useful starting point for developing flight models for newly invaded areas.

Slow-growth high-mortality: A meta-analysis for insects.

The slow growth-high mortality hypothesis (SG-HG) predicts that slower growing herbivores suffer greater mortality due to a prolonged window of vulnerability. Given diverse plant-herbivore-natural enemy systems resulting from different feeding ecologies of herbivores and natural enemies, this hypothesis might not always be applicable to all systems. This is evidenced by mixed support from empirical data. In this study, a meta-analysis of the SG-HM hypothesis for insects was conducted, aiming to find conditions that favor or reject SG-HM. The analysis revealed significant within- and between-group heterogeneity for almost all explanatory variables and overall did not support SG-HM. In this analysis, SG-HM was supported when any of the following 5 conditions was met: (1) host food consisted of artificial diet; (2) herbivore growth was measured as larval mass; (3) herbivores were generalists; (4) no or multiple species of natural enemies were involved in the study; and (5) parasitoids (i.e., parasitic insects) involved in the study were gregarious. SG-HM was rejected when any of the following 5 conditions was met: (1) herbivores were from the order Hymentoptera; (2) parasitoids from more than 1 order caused herbivore mortality; (2) parasitoids were specialists; (3) parasitoids were solitary; (4) parasitoids were idiobionts or koinobionts; and (5) single species of natural enemy caused mortality of specialist herbivores. All known studies investigated herbivore mortality for a short period of their life cycle. Researchers are encouraged to monitor herbivore mortality during the entire window of susceptibility or life cycle using life tables. Studies involving multiple mortality factors (i.e., both biotic and abiotic) or multiple natural enemy species are also encouraged since herbivores in nature face a multitude of risks during the entire life cycle. More comprehensive studies may increase our understanding of factors influencing the relationships between herbivore growth and mortality.

Comparative morphometric and chemical analyses of phenotypes of two invasive ambrosia beetles (Euwallacea spp.) in the United States.

The polyphagous shot hole borer (PSHB), Euwallacea sp., was first detected in 2003 in Los Angeles County, California, USA. Recently, this invasive species has become a major pest of many hardwood trees in urban and wildland forests throughout southern California. PSHB is nearly identical in morphology and life history to the tea shot hole borer (TSHB), Euwallacea fornicatus, an invasive pest of hardwoods in Florida, USA and many other parts of the world. However, molecular studies have suggested that the taxa are different species. We conducted morphometric and chemical analyses of the phenotypes of Euwallacea sp. collected in southern California (Los Angeles County) and E. fornicatus collected in Florida (Miami-Dade County). Our analyses indicated that PSHB has 3 larval instars. The third larval instar was separated from the first 2 instars by head capsule width with 0 probability of misclassification. The body length, head width, and pronotal width of PSHB adult males were significantly less than those of females. Head width and pronotal width of female PSHB were significantly less than those of female TSHB. In contrast, body length, and ratio of body length to pronotal width of female PSHB were significantly greater than those of female TSHB. However, females of these 2 species could not be separated completely by these 4 measurements because of the overlapping ranges. Cuticular hydrocarbons detected in both species were exclusively alkanes (i.e., n-alkanes, monomethylalkanes, dimethylalkanes, and trimethylalkanes). Cuticular hydrocarbon profiles of PSHB males and females were similar, but they both differed from that of TSHB females. Cuticular hydrocarbons of PSHB were predominantly internally branched dimethylalkanes with backbones of 31 and 33 carbons, whereas cuticular hydrocarbons of TSHB females were dominated by internally branched monomethylalkanes and dimethylalkanes with backbones of 28 and 29 carbons. Multiple compounds within these classes appear to be diagnostic for PSHB and TSHB, respectively.

Abundance of volatile organic compounds in white ash phloem and emerald ash borer larval frass does not attract Tetrastichus planipennisi in a Y-tube olfactometer.

Many natural enemies employ plant- and/or herbivore-derived signals for host/prey location. The larval parasitoid Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) is 1 of 3 biocontrol agents currently being released in an effort to control the emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coloeptera: Burprestidae) in North America. To enhance its efficiency, allelochemicals that attract it need to be assessed. In this study, ash phloem volatile organic compounds (VOCs) of black, green, and white ash, and EAB larval frass were compared. Foraging behavior of T. planipennisi females in response to VOCs of white ash or frass from EAB larvae feeding on white ash phloem was tested using a Y-tube olfactometer. Results indicated that the 3 ash species had similar VOC profiles. EAB larval frass generally contained greater levels of VOCs than phloem. Factor analysis indicated that the 11 VOCs could be broadly divided into 2 groups, with α-bisabolol, ß-caryophyllene, (E)-2-hexenal, (Z)-3-hexenal, limonene, methyl benzoate, methyl indole-3-acetic acid, methyl jasmonate, methyl salicylate as the first group and the rest (i.e., methyl linoleate and methyl linolenate) as a second. Abundance of VOCs in white ash phloem tissue and frass, nevertheless, did not attract T. planipennisi females. The concealed feeding of EAB larvae might explain the selection for detectable and reliable virbrational signals, instead of undetectable and relatively unreliable VOC cues from phloem and frass, in short-range foraging by T. planipennisi. Alternatively, it is possible that T. planipennisi is not amenable to the Y-tube olfactometer assay employed.

Impact of the goldspotted oak borer, Agrilus auroguttatus, on the health of coast live oak before and after treatment with two systemic insecticides.

BACKGROUND: The invasive goldspotted oak borer, Agrilus auroguttatus, is threatening the health and survival of oak trees in San Diego County, California. From two sites in the core area of the infestation, we report a 2.5 year investigation of the impact of A. auroguttatus on coast live oak, Quercus agrifolia, before and after treatment with two systemic insecticides, emamectin benzoate (EB) and imidacloprid (IC). RESULTS: None of the 446 survey trees died during the study. The crown dieback rating of most trees at both study sites remained unchanged, regardless of insecticide treatment. A higher cumulative increase in the number of A. auroguttatus emergence holes was observed on trees that were previously infested and on trees with larger diameters. Over the 2.5 year period, the new infestation rates of initially uninfested trees across the untreated and treated groups were 50% (EB) and 32% (IC), and neither EB nor IC treatment affected cumulative increases in the number of emergence holes. EB-injected trees did not have significant annual increases in the number of A. auroguttatus emergence holes at either 1.5 or 2.5 years compared with that at 0.5 years, whereas untreated trees had significant annual increases. Although IC-injected trees had a significantly greater annual increment in the number of emergence holes than untreated trees during the last year of the study, treated trees had significant reductions in annual increases in emergence holes at both 1.5 and 2.5 years compared with that at 0.5 years. Untreated trees had no significant reduction in the annual increase in emergence holes at 1.5 and 2.5 years. CONCLUSIONS: A. auroguttatus preferentially attacked previously infested and larger (diameter at breast height > 15-30 cm) oak trees, but the attacks led to very gradual changes in the health of the trees. Both EB and IC provided minor suppressive effects on A. auroguttatus emergence. Published 2014. This article is a U.S. Government work and is in the public domain in the USA.

Crepuscular flight activity of an invasive insect governed by interacting abiotic factors.

Seasonal and diurnal flight patterns of the invasive walnut twig beetle, Pityophthorus juglandis, were assessed between 2011 and 2014 in northern California, USA in the context of the effects of ambient temperature, light intensity, wind speed, and barometric pressure. Pityophthorus juglandis generally initiated flight in late January and continued until late November. This seasonal flight could be divided approximately into three phases (emergence: January-March; primary flight: May-July; and secondary flight: September-October). The seasonal flight response to the male-produced aggregation pheromone was consistently female-biased (mean of 58.9% females). Diurnal flight followed a bimodal pattern with a minor peak in mid-morning and a major peak at dusk (76.4% caught between 1800 and 2200 h). The primarily crepuscular flight activity had a Gaussian relationship with ambient temperature and barometric pressure but a negative exponential relationship with increasing light intensity and wind speed. A model selection procedure indicated that the four abiotic factors collectively and interactively governed P. juglandis diurnal flight. For both sexes, flight peaked under the following second-order interactions among the factors when: 1) temperature between was 25 and 30 °C and light intensity was less than 2000 lux; 2) temperature was between 25 and 35 °C and barometric pressure was between 752 and 762 mba (and declined otherwise); 3) barometric pressure was between 755 and 761 mba and light intensity was less than 2000 lux (and declined otherwise); and 4) temperature was ca. 30 °C and wind speed was ca. 2 km/h. Thus, crepuscular flight activity of this insect can be best explained by the coincidence of moderately high temperature, low light intensity, moderate wind speed, and low to moderate barometric pressure. The new knowledge provides physical and temporal guidelines for the application of semiochemical-based control techniques as part of an IPM program for this invasive pest.

Developing monitoring techniques for the invasive goldspotted oak borer (Coleoptera: Buprestidae) in California.

The goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), is an invasive species that has colonized oak woodlands in southern California. To better define its seasonal flight activity, assist with forest and integrated pest management activities, and define the current distribution in California, an effective monitoring technique for A. auroguttatus is necessary. We assessed the efficacy of two colors of flight-intercept prism traps, the placement of these traps at three heights, and several commercially available lures [Manuka oil, Phoebe oil, and a green leaf volatile, (3Z)-hexenol] for monitoring the flight of adult A. auroguttatus. Landing rates and the densities of D-shaped emergence holes of A. auroguttatus adults were assessed on the lower stems of coast live oak, Quercus agrifolia Née, of varying size and crown health classes. Purple flight-intercept prism traps placed at heights of 3 m and 4.5 m caught significantly more female A. auroguttatus than green prism traps. In one experiment, males also responded at a significantly higher level to purple than to green prism traps placed at 3 m height. The addition of commercially available lures significantly enhanced male, but not female, A. auroguttatus trap catch when compared with unbaited control traps. There were no differences among male flight responses to the three lures. A. auroguttatus landing rates and emergence hole densities were significantly greater on the largest-diameter trees (>76.2 cm diameter at breast height) and on trees with severe crown thinning or complete crown collapse. The annual increment in emergence hole densities was also significantly greater on trees with severe crown thinning or complete crown collapse. In three trapping studies over multiple years in southern California, the adult flight period began as early as mid-May, peaked in mid-June to early July, and ended in early- to mid-September. To demonstrate the efficacy of the detection method for A. auroguttatus (unbaited purple traps at 3 m height), a delimitation survey conducted from 2009 to 2012 confirmed that the species was only present in San Diego Co., but that the distribution was expanding northward.

Analyzing blends of herbivore-induced volatile organic compounds with factor analysis: revisiting "cotton plant, Gossypium hirsutum L., defense in response to nitrogen fertilization".

Many herbivorous, predaceous, and parasitic insects use constitutive and herbivore-induced volatile organic compounds (VOCs) to locate their respective host plant, prey, and hosts. Multivariate statistical tools (e.g., factor analysis) are recognized increasingly as an appropriate approach for analyzing intercorrelated data such as presence/absence or quantities of VOCs. One challenge of implementing factor analysis is determining how many new variables (factors) to retain in the final analysis. I demonstrate a method proposed by Johnson and Wichern to mitigate this problem by using VOC data published in Chen et al. The advantage of using loading (or weight) transformation in interpretation of new variables was also illustrated in the example. Factor analysis found similar nitrogen fertilization effects on VOC production as those in Chen et al. Similarities were 1) nitrogen fertilization interacted with herbivore damage status on VOC production: at low nitrogen (42 ppm) level, beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), damage elicited increases in VOC production, whereas at high nitrogen (196 ppm) VOC production was suppressed; 2) nitrogen fertilization did not affect limonene, alpha-pinene, and beta-pinene production. The seven individual VOCs significantly affected by nitrogen fertilization in Chen et al. were (Z)-3-hexenal, (E)-2-hexenal, (E)-beta-farnesene, (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), alpha-bergamotene, gamma-bisabolene, and bisabolol, of which only three ((E)-beta-farnesene, gamma-bisabolene, and bisabolol) weighed heavily on factor 1 in the current study.

Feeding by emerald ash borer larvae induces systemic changes in black ash foliar chemistry.

The exotic wood-boring pest, emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), has been threatening North American ash (Fraxinus spp.) resources, this being recognized since its first detection in Michigan, USA and Ontario, Canada in 2002. Ash trees are killed by larval feeding in the cambial region, which results in disruption of photosynthate and nutrient translocation. In this study, changes in volatile and non-volatile foliar phytochemicals of potted 2-yr-old black ash, Fraxinus nigra Marshall, seedlings were observed in response to EAB larval feeding in the main stem. EAB larval feeding affected levels of six compounds [hexanal, (E)-2-hexenal, (Z)-3-hexenyl acetate, (E)-ß-ocimene, methyl salicylate, and (Z,E)-α-farnesene] with patterns of interaction depending upon compounds of interest and time of observation. Increased methyl salicylate emission suggests similarity in responses induced by EAB larval feeding and other phloem-feeding herbivores. Overall, EAB larval feeding suppressed (Z)-3-hexenyl acetate emission, elevated (E)-ß-ocimene emission in the first 30days, but emissions leveled off thereafter, and generally increased the emission of (Z,E)-α-farnesene. Levels of carbohydrates and phenolics increased overall, while levels of proteins and most amino acids decreased in response to larval feeding. Twenty-three amino acids were consistently detected in the foliage of black ash. The three most abundant amino acids were aspartic acid, glutamic acid, glutamine, while the four least abundant were α-aminobutyric acid, ß-aminoisobutyric acid, methionine, and sarcosine. Most (16) foliar free amino acids and 6 of the 9 detected essential amino acids decreased with EAB larval feeding. The ecological consequences of these dynamic phytochemical changes on herbivores harbored by ash trees and potential natural enemies of these herbivores are discussed.

Role of emerald ash borer (Coleoptera: Buprestidae) larval vibrations in host-quality assessment by Tetrastichus planipennisi (Hymenoptera: Eulophidae).

The biological control agent Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) is a gregarious larval endoparasitoid of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive cambium-feeding species responsible for recent, widespread mortality of ash (Fraxinus spp.) in North America. T. planipennisi is known to prefer late-instar emerald ash borer, but the cues used to assess host size by this species and most other parasitoids of concealed hosts remain unknown. We sought to test whether vibrations produced by feeding emerald ash borer vary with larval size and whether there are any correlations between these cues and T. planipennisi progeny number (i.e., brood size) and sex ratio. The amplitudes and rates of 3-30-ms vibrational impulses produced by emerald ash borer larvae of various sizes were measured in the laboratory before presenting the larvae to T. planipennisi. Impulse-rate did not vary with emerald ash borer size, but vibration amplitude was significantly higher for large larvae than for small larvae. T. planipennisi produced a significantly higher proportion of female offspring from large hosts than small hosts and was shown in previous work to produce more offspring overall from large hosts. There were no significant correlations, however, between the T. planipennisi progeny data and the emerald ash borer sound data. Because vibration amplitude varied significantly with host size, however, we are unable to entirely reject the hypothesis that T. planipennisi and possibly other parasitoids of concealed hosts use vibrational cues to assess host quality, particularly given the low explanatory potential of other external cues. Internal chemical cues also may be important.

Differential response in foliar chemistry of three ash species to emerald ash borer adult feeding.

The emerald ash borer (EAB; Agrilus planipennis Fairmaire; Coleoptera: Buprestidae), is an exotic wood-boring beetle that has been threatening North American ash (Fraxinus spp.) resources since its discovery in Michigan and Ontario in 2002. In this study, we investigated the phytochemical responses of the three most common North American ash species (black, green, and white ash) in northeastern USA to EAB adult feeding. Black ash was the least responsive to EAB adult feeding in terms of the induction of volatile compounds, and levels of only two (indole and benzyl cyanide) of the 11 compounds studied increased. In green ash, levels of two [(E)-ß-ocimene and indole] of the 11 volatile compounds studied were elevated, while the levels of two green leaf volatiles [hexanal and (E)-2-hexenal] decreased. White ash showed the greatest response with an increase in levels of seven of the 11 compounds studied. Qualitative differences among ash species were detected. Among the phenolic compounds detected, ligustroside was the only one detected in all three species. Oleuropein aglycone and 2 unidentified compounds were found only in black ash; coumaroylquinic acid and feruloylquinic acid were detected only in green ash; and verbascoside hexoside was detected only in white ash. EAB adult feeding did not elicit or decrease concentrations of any selected individual phenolic compounds. However, although levels of total phenolics from black and green ash foliage were not affected by EAB adult feeding, they decreased significantly in white ash. EAB adult feeding elevated chymotrypsin inhibitors in black ash. The possible ecological implications of these findings are discussed.

Biotic and abiotic factors affect green ash volatile production and emerald ash borer adult feeding preference.

The emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is an exotic woodborer first detected in 2002 in Michigan and Ontario and is threatening the ash resource in North America. We examined the effects of light exposure and girdling on green ash (Fraxinus pennsylvanica Marsh) volatile production, and effects of light exposure, girdling, and leaf age on emerald ash borer adult feeding preferences and phototaxis. Green ash seedlings grown under higher light exposure had lower amounts of three individual volatile compounds, (Z)-3-hexenol, (E)-beta-ocimene, and (Z,E)-alpha-farnesene, as well as the total amount of six detected volatile compounds. Girdling did not affect the levels of these volatiles. Emerald ash borer females preferred mature leaves, leaves from girdled trees, and leaves grown in the sun over young leaves, leaves from nongirdled trees, and leaves grown in the shade, respectively. These emerald ash borer preferences were most likely because of physical, nutritional, or biochemical changes in leaves in response to the different treatments. Emerald ash borer females and males showed positive phototaxis in laboratory arenas, a response consistent with emerald ash borer preference for host trees growing in sunlight.

Interactive influence of leaf age, light intensity, and girdling on green ash foliar chemistry and emerald ash borer development.

Biotic and abiotic environmental factors affect plant nutritional quality and defensive compounds that confer plant resistance to herbivory. Influence of leaf age, light availability, and girdling on foliar nutrition and defense of green ash (Fraxinus pennsylvanica Marsh) was examined in this study. Longevity of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), adults reared on green ash foliage subjected to these factors was assayed. Mature leaves generally were more nutritious with greater amino acids and a greater ratio of protein to non-structural carbohydrate (P:C) than young leaves, in particular when trees were grown in shade. On the other hand, mature leaves had lower amounts of trypsin and chymotrypsin inhibitors, and total phenolics compared to young leaves. Lower defense of mature leaves alone, or along with higher nutritional quality may lead to increased survival and longevity of emerald ash borer feeding on mature leaves. Sunlight reduced amino acids and P:C ratio, irrespective of leaf age and girdling, and elevated total protein of young foliage, but not protein of mature leaves. Sunlight also dramatically increased all investigated defensive compounds of young, but not mature leaves. Girdling reduced green ash foliar nutrition, especially, of young leaves grown in shade and of mature leaves grown in sun. However emerald ash borer performance did not differ when fed leaves from trees grown in sun or shade, or from girdled or control trees. One explanation is that emerald ash borer reared on lower nutritional quality food may compensate for nutrient deficiency by increasing its consumption rate. The strong interactions among leaf age, light intensity, and girdling on nutrition and defense highlight the need for caution when interpreting data without considering possible interactions.

Changes of oxidase and hydrolase activities in pecan leaves elicited by black pecan aphid (Hemiptera: Aphididae) feeding.

The black pecan aphid, Melanocallis caryaefoliae (Davis) (Hemiptera: Aphididae), is a foliar feeder of pecan, Carya illinoinensis (Wangenh.) K. Koch (Juglandaceae). The pest causes chlorosis of leaflet lamina, physiological damage to foliage and trees, and commonly limits the profitability of commercial pecan orchard enterprises. However, key aspects of this host-pest interaction are poorly understood. We report here the effects of M. caryaefoliae feeding on the foliar activity of oxidative (i.e., catalase, lipoxygenase [LOX]-1 and 3, and peroxidase) and hydrolytic (i.e., esterase) enzymes in relation to the degree of aphid resistance among pecan varieties. The 2-yr study showed that M. caryaefoliae-infested foliage exhibited elevated peroxidase activity only in susceptible ('Desirable', 'Sumner', and 'Schley'), but not in resistant ('Cape Fear', 'Gloria Grande', and 'Money Maker') genotypes. Susceptible genotypes also exhibited more severe leaf chlorosis in response to M. caryaefoliae feeding than the resistant genotypes; however, the aphid feeding did not influence catalase or esterase activity in all varieties, except the increase of esterase activity in Desirable and Gloria Grande. Melanocallis caryaefoliae feeding also influences activity of two lipoxygenase isozymes, with LOX3 being more frequently induced than LOX1. Foliar LOX3 activity was more frequently induced by M. caryaefoliae feeding in the moderately resistant 'Oconee' and highly resistant Money Maker and Cape Fear than in the susceptible genotypes. Therefore, the elevation of peroxidase is likely to be associated with aphid susceptibility and contributed to the severe leaf chlorosis, whereas the increase of LOX3 activity might be associated with aphid resistance in pecan. These findings contribute to our understanding of the etiology of M. caryaefoliae-elicited leaf chlorosis on pecan foliage. Such information may also be used to develop enzyme markers for identifying black pecan aphid resistance and/or susceptibility in pecan germplasm.

Physiological, nutritional, and biochemical bases of corn resistance to foliage-feeding fall armyworm.

Three corn (Zea mays) germplasm lines [i.e., Ab24E (susceptible control), Mp708 (resistant control), and a locally selected partial inbred line FAW7050 (resistant)] were examined for Spodoptera frugiperda (J.E. Smith; Lepidoptera: Noctuidae) resistance. Nutritional [i.e., total protein content, amino acids, glucose, total nonstructural carbohydrates (TNC), protein to TNC (P/C) ratios] and biochemical (i.e., peroxidase and lipoxygenase 3) properties in the seedlings of these corn lines were examined to categorize resistance mechanisms to S. frugiperda. Physiological changes in photosynthetic rates also were examined in an attempt to explain nutritional and biochemical dynamics among corn germplasm lines and between insect-infested and noninfested corn plants within a germplasm line. Results indicated that S. frugiperda larvae survived better and developed faster in susceptible Ab24E than in resistant FAW7050 or Mp708. The three germplasm lines differed in resistance mechanisms to S. frugiperda, and the observed patterns of resistance were probably collective results of the P/C ratio and defensive proteins. That is, the susceptibility of Ab24E to S. frugiperda was due to a high P/C ratio and a low level of induced defensive compounds in response to insect herbivory, while the resistance of FAW7050 resulted from elevated defensive proteins following insect herbivory, low P/C ratio, and elevated defensive proteins in Mp708 contributed to its resistance to S. frugiperda. The elevated protein amounts in resistant Mp708 and FAW7050 following S. frugiperda injury were likely due to greater conversion of photosynthates to defensive proteins following the greater photosynthetic rates in these entries. Greater photosynthetic capacity in Mp708 and FAW7050 also led to higher amino acid and glucose contents in these two lines. Neither amino acid nor lipoxygenase 3 played a critical role in corn resistance to S. frugiperda. However, high inducibility of peroxidase may be an indicator of S. frugiperda susceptibility as observed elsewhere.

Cotton plant, Gossypium hirsutum L., defense in response to nitrogen fertilization.

Plants respond to insect herbivory by producing dynamic changes in an array of defense-related volatile and nonvolatile secondary metabolites. A scaled response relative to herbivory levels and nutrient availability would be adaptive, particularly under nutrient-limited conditions, in minimizing the costs of expressed defensive pathways and synthesis. In this study, we investigated effects of varying nitrogen (N) fertilization (42, 112, 196, and 280 ppm N) on levels of cotton plant (Gossypium hirsutum) phytohormones [jasmonic acid (JA) and salicylic acid (SA)], terpenoid aldehydes (hemigossypolone, heliocides H(1), H(2), H(3), and H(4)), and volatile production in response to beet armyworm (Spodoptera exigua) herbivory. Additional bioassays assessed parasitoid (Cotesia marginiventris) host-searching success in response to cotton plants grown under various N fertilizer regimes. At low N input (42 ppm N), herbivore damage resulted in significant increases in local leaf tissue concentrations of JA and volatiles and in systemic accumulation of terpenoid aldehydes. However, increased N fertilization of cotton plants suppressed S. exigua-induced plant hormones and led to reduced production of various terpenoid aldehydes in damaged mature leaves and undamaged young leaves. While increased N fertilization significantly diminished herbivore-induced leaf volatile concentrations, the parasitism of S. exigua larvae by the parasitoid C. marginiventris in field cages did not differ among N treatments. This suggests that, despite significant N fertilization effects on herbivore-induced plant defenses, at short range, the parasitoids were unable to differentiate between S. exigua larvae feeding on physiologically different cotton plants that share large constitutive volatile pools releasable when damaged by herbivores.

[Diversity and stability of arthropod assemblage in tea orchard].

Two tea orchards, simplex tea orchard with weeds removed manually or by herbicides (STO) and complex tea orchard with the weed Hedyotis uncinella (CTO), each with an area of 0. 4 hm2, were established in 1995 in Yingde Hongxing Tea Plantation, Guangdong Province. The primary eigenvalues, species richness index (R), assemblage diversity index (H'), evenness index (J) and species concentration index (C) of arthropod assemblage were employed and compared to assess the efficacy of STO and CTO on the diversity and stabilityof arthropod assemblage. Stability indexes Ss/Si and Sn/Sp and variation coefficient of diversity index ds/dm were utilized as well. The results demonstrated that the R of arthropod assemblage in CTO ranged from 4 to 8, with the highest of 7.7403, while that in STO varied mainly between 4 to 6. The average R of arthropod assemblage in CTO was 5.4672 +/- 0.3483, higher than that in STO (4.8809 +/- 0.3175). The H' of arthropod in CTO (3.8535 +/- 0.1232) was higher, in contrast to the value in STO (3.4654 +/- 0.1856). The J in CTO was higher, while the species concentration index (C) was lower, in comparison to STO. The stability indexes Ss/Si and Sn/Sp of CTO were greater than those of STO, while the ds/dm in CTO (0.1107) was lower than that in STO (0.1855). All these indicated that the diversity of arthropod assemblage was better preserved in CTO, and the assemblage in CTO was more stable.