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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 walnut twig beetle, Pityophthorus juglandis Blackman (Coleoptera: Scolytidae), vectors a phytopathogenic fungus, Geosmithia morbida Kolařik et al. (Hypocreales), which causes thousand cankers disease (TCD) in walnut (Juglans sp.) and wingnut (Pterocarya sp., both Juglandaceae) trees. We investigated an early point in disease inception in two walnut species – Juglans californica S. Wats. and Juglans major (Torr. ex Sitsgr.) Heller – native to riparian forests of the western USA by comparing P. juglandis flight and landing responses to small‐diameter branch sections. Twenty unbaited branch sections (10 each of J. californica and J. major) were presented in a completely randomized design to populations of P. juglandis at the USDA Agricultural Research Service, National Clonal Germplasm Repository (NCGR) Juglans collection located at Wolfskill Experimental Orchards (Winters, CA, USA) and at the California State University, Chico, Agricultural Teaching and Research Center (ATRC, Chico, CA). These assays were carried out within a 4‐ to 6‐year period when weekly flight surveys with aggregation pheromone‐baited multiple funnel traps revealed that P. juglandis flight activity–abundance was higher at the NCGR than at the ATRC. For the landing rate assays, adhesive‐coated acetate sheets were wrapped around the branch sections and exchanged weekly. Three assays were completed at the NCGR (assays 1–3), whereas one assay was completed at the ATRC (assay 4). Landing rates on these traps were compared between J. californica and J. major. Two additional assays (5 and 6) were completed at the NCGR to compare responses to branch sections of J. californica and to similarly sized cardboard tubes (negative control). All six assays were completed over a 4‐year span during the 4‐ to 6‐year weekly flight survey period. Pooled landing rates of male and female P. juglandis (assays 1–4) demonstrated a preference by both sexes for J. californica over J. major. In assay 5 there was no preference by males or females for J. californica over the negative control, perhaps due to the low flight activity–abundance of P. juglandis during the assay. When repeated at a time of higher flight activity–abundance (assay 6), male and female landing rates on J. californica exceeded those on the negative control. Females of the invasive fruit‐tree pinhole borer (an ambrosia beetle), Xyleborinus saxeseni (Ratzeburg), and an invasive bark beetle, Hypothenemus eruditus Westwood (both Coleoptera: Scolytidae), showed relatively higher flight responses than either sex of P. juglandis during most assays, suggesting higher population densities of these two other invasive species at the two orchards or a greater sensitivity to host volatiles. Xyleborinus saxeseni and H. eruditus preferred to land on J. major over J. californica and on J. californica over the negative control. Similarly, an invasive longhorned beetle, Nathrius brevipennis (Mulsant) (Coleoptera: Cerambycidae), showed a significant preference for J. major over J. californica, but not for J. californica over the negative control. More male N. brevipennis were trapped than females at both study sites [sex ratio ranged from 5:1 (assay 6) to 39:1 (assay 4)], and flight occurred only in the spring and early summer months. Another ambrosia beetle trapped at the NCGR and ATRC, Xyleborus affinis Eichhoff, represented the first records of this species from western North America. In summary, flight responses recorded in some of our assays for P. juglandis and several other subcortical insects on Juglans indicate that host preference by these insects may be determined by long‐range olfactory cues that do not involve pheromones.

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.

The predatory stink bug Eocanthecona furcellata (Wolff) (Hemiptera: Pentatomidae), which is widely distributed in the subtropics and tropics, is a potential biological control agent for lepidopteran, coleopteran, and hemipteran pests. We investigated development of eggs oviposited by E. furcellata females subjected to various photoperiods (i.e., 8L:16D, 12L:12D, and 16L:8D) and temperature regimes (i.e., 26:26, 30:28, 32:30, 34:32, 35:33, and 36:34 °C; temperatures at light and dark phases, respectively) at different developmental stages. Exposure of E. furcellata nymphs and adults to different photoperiods and temperature regimes affected the development of eggs they subsequently oviposited. Temperatures greater than 30 °C induced diapause in a portion of the eggs, as evidenced by the delayed appearance of the black circle and the red spot, characteristics of early and late stage of embryonic development, respectively. The proportion of diapausing eggs generally increased with increasing temperature exposure, whereas it decreased with advancing developmental stage/age at which E. furcellata was subjected to the temperature regime. Although viability of diapausing eggs was lower than that of non-diapausing eggs, diapausing eggs survived longer under high temperatures. The trade-off between survivability and viability may be a strategy to survive unpredictability of summer temperatures.

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.

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.

We assessed the impact of the polyphagous shot hole borer (PSHB), Euwallacea whitfordiodendrus (Schedl), and Kuroshio shot hole borer (KSHB), E. kuroshio Gomez and Hulcr, on hardwood trees in southern California, southwestern China, and northwestern Vietnam. The highest levels of mortality were recorded from 10 of 39 tree species in the survey, and these were primarily native tree species. Two invasive shot hole borers represent relatively recent introductions in southern California, USA, and continue to spread and cause injury and mortality to several native and ornamental tree species. They originate from Southeast Asia. Knowledge of tree species susceptibility to these wood-boring beetles is essential to inform better pest management and to evaluate future risk for urban and wildland forests. From 2012 to 2016, ground surveys were conducted in the invaded and native regions at PSHB/KSHB-infested and PSHB/KSHB-uninfested sites to record levels of tree injury and mortality on native and ornamental tree species. In California, several native species of maple, Acer, willow, Salix, and sycamore, Platanus, were infested by either PSHB or KSHB at high rates (> 70%), and comparative rate of infestation by KSHB in all trees and in native trees surpased that by PSHB, whereas rate of infestation by PSHB in ornamental trees surpassed that by KSHB. Mortality of two maple species caused by PSHB exceeded 20%, whereas background mortality rate of hardwoods was 2% in uninfested areas. These data should inform land managers about the tree species at most risk to injury and mortality, facilitate detection ground surveys, and direct prophylactic treatments for these invasive woodborers.

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, β-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.

As of summer 2014, the invasive emerald ash borer (EAB),Agrilus planipennisFairmaire (Coleoptera: Buprestidae), has become established in 24 states in the United States of America and has killed tens of millions of ash trees since its introduction into Michigan in the 1990s. Considerable research has been conducted on many aspects of EAB life history, natural history, ecology, and management strategies in an attempt to contain this devastating pest. In this article, we review the life history, mating behaviours, and host plant selection by EAB in North America as well as host resistance to EAB attack.

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.

Nitrogen has complex effects on plant–herbivore–parasitoid tritrophic interactions. The negative effects of low nitrogen fertilization in host plants on insect herbivores can be amplified to the higher trophic levels. In the present study, we examined the impact of varying nitrogen fertilization (42, 112, 196, and 280 ppm) of cotton plants ( Gossypium hirsutum L.) on the interactions between the beet armyworm, Spodoptera exigua (Hubner) (Lepidoptera: Noctuidae), and the hymenopteran endoparasitoid Cotesia marginiventris (Cresson) (Hymenoptera: Braconidae). We predicted that the development and fitness of C. marginiventris would be adversely affected by low host plant nitrogen fertilization through the herbivore S. exigua . The percentage of C. marginiventris offspring developing to emerge and spin a cocoon, and total mortality of parasitized S. exigua larvae were unaffected by nitrogen level. The developmental time of C. marginiventris larvae in S. exigua larvae feeding on low (42 ppm) nitrogen cotton plants was approximately 30% longer than that of those feeding on higher (112, 196, and 280 ppm) nitrogen plants. Parasitoid size (length of right metathoracic tibia), a proxy for fitness, of C. marginiventris males was positively affected by nitrogen level. Total amounts of S. exigua hemolymph proteins were not affected by nitrogen level, but were reduced by parasitism by C. marginiventris . Two proteins with molecular weights of ca. 84 and 170 kDa dominated the S. exigua larval hemolymph proteins. Concentrations of the 170 kDa hemolymph protein were unaffected by nitrogen treatment, but parasitism reduced concentrations of the 170 kDa protein. Concentrations of the 84 kDa protein, on the other hand, were interactively affected by parasitism and nitrogen treatment: higher nitrogen fertilization (112, 196, and 280 ppm) increased protein concentrations relative to the 42 ppm treatment for unparasitized S. exigua larvae, whereas nitrogen treatment had no effects on parasitized larvae. For S. exigua larvae feeding on 42 ppm nitrogen plants, parasitism increased concentration of the 84 kDa protein, while for those feeding on 112, 196, and 280 ppm nitrogen plants, parasitism decreased concentrations of the protein. Possible mechanisms and ecological consequences for the extended development of C. marginiventris on S. exigua hosts grown on low-nitrogen plants are discussed.

Adults of the invasive goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), consumed foliar weight in no-choice feeding tests of, in descending order, California black oak Quercus kelloggii Newb., Engelmann oak, Quercus engelmannii Greene, coast live oak, Quercus agrifolia Nee, and canyon live oak, Quercus chrysolepis Liebm. (Fagaceae). Furthermore, significantly more foliar area was consumed of Q. kelloggii than of Q. chrysolepis. In dual-choice feeding tests with isolated leaf disks, A. auroguttatus consumed significantly more foliar weight and area of Q. kelloggii relative to the other three oak species, and more foliar weight of Q. agrifolia than of Q. chrysolepis. In dual-choice feeding tests with leaves on small branches, A. auroguttatus consumed more foliar weight of Q. kelloggii than of Q. engelmannii and Q. agrifolia. Thus, multiple experiments suggested that adults of A. auroguttatus preferred the foliage of Q. kelloggii over that of the other three oak species, and among the other three species they did not appear to have a strong feeding preference. Factor analysis reduced the quantities of 13 foliar nutrients into two new variables (factor 1 and factor 2). Factor 1 was weighted heavily on the quantities of nitrogen, sulfur, phosphorus, potassium, zinc, and copper, whereas factor 2 was weighted heavily on the quantities of zinc, iron, and aluminum. Factor 1 varied by oak species, with Q. kelloggii having a higher factor 1 nutrient content than the other three species. Factor 2 response was higher in Q. kelloggii, Q. agrifolia, and Q. engelmannii than in Q. chrysolepis. The collective effects of four macronutrients (nitrogen, sulfur, phosphorus, and potassium) and two micronutrients (zinc and copper) suggest that these might be the nutrients directing preferential feeding of A. auroguttatus adults on the foliage of Q. kelloggii. Leaf toughness might also play an important role in feeding preference. Female A. auroguttatus did not show an ovipositional preference among the four oak species.

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.

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.

From 2009–2011, we assessed the impact of the goldspotted oak borer, Agrilus auroguttatus Schaeffer, or its sibling species, Agrilus coxalis Waterhouse, at locations in southern California (denoted infested: ICA and uninfested: UCA), southeastern Arizona (AZ), and southern Mexico (MX). Our surveys examined forest composition of oak woodlands; the degree of injury and proportion of oaks infested with either A. auroguttatus (ICA and AZ) or A. coxalis (MX); and the progression of aerially mapped oak mortality in San Diego Co. (ICA). By most measures of impact that we evaluated, the effect on oaks by the two Agrilus spp. was relatively low in their native regions, but significantly higher by A. auroguttatus at ICA sites. Larger diameter red oak species have been the preferred hosts of A. auroguttatus in AZ and ICA sites, and red oaks greater than approx. 13 cm in DBH throughout California are likely at risk of injury from this invasive pest. At sites in AZ there was no evidence of infestation by A. auroguttatus on living or dead white oak species, whereas at ICA sites we recorded a minor amount of infestation by A. auroguttatus on living individuals of a white oak species (Quercus engelmannii Greene), but no mortality. In contrast, a white oak from MX sites (Quercus peduncularis) was more frequently infested by A. coxalis than were indigenous red oaks. Across all ICA sites, A. auroguttatus has infested 61% of the live larger diameter oaks and killed 13% of the oak component of the forest (vs. 4% infested and 2% dead in AZ, respectively). At survey plots near the predicted origin of the outbreak in CA, over 90% of the larger diameter red oaks have been infested. Nearly 90% of the dead oaks surveyed across all ICA sites showed evidence of previous injury symptoms from A. auroguttatus. Aerial oak mortality polygons associated with A. auroguttatus have expanded � 50 km in nine years, but our analysis confirms that the outbreak appears to still be confined to San Diego Co. The distance of oak mortality polygons from the predicted origin of the outbreak explained the most variance in a principal component analysis. The invasive population of A. auroguttatus is a significant conservation and ecological threat to the oak woodlands of California and should be managed accordingly, especially by restricting firewood movement. Published by Elsevier B.V.

Two experiments were performed to determine the extent to which ash species (black, green and white) and larval developmental stage (second, third and fourth instar) affect the efficiency of phloem amino acid utilization by emerald ash borer (EAB) Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) larvae. EAB larvae generally utilized green ash amino acids more efficiently than those of the other two species. For example, the concentrations of only six (two essential) and seven (two essential) amino acids were lower in frass from EAB that fed upon black and white ash than in the corresponding phloem, respectively. By contrast, concentrations of 16 (eight essential) amino acids were lower in the frass from EAB that fed upon green ash than in the phloem. In addition, in green ash, the frass : phloem ratios of 13 amino acids were lower than their counterparts in black and white ash.

The diurnal flight response of the walnut twig beetle, Pityophthorus juglandis Blackman (Coleoptera: Scolytidae), was assessed during two seasonal periods at two sites in northern California. Males and females flew primarily at dusk in response to aggregation pheromone-baited traps during late June/early July, and the percentage of beetles that flew between sunrise and late afternoon was positively correlated with the temperature at 6 AM of that day. Between late August and early November, the diurnal flight pattern was more varied and generally bimodal, though approx. 42 to 55% of the beetles still flew during the dusk period. During either period of the season, there was very little evidence that any flight occurred during the night. Between late August and late September, diurnal flight of both sexes during two-hour intervals was associated with temperature in a Gaussian manner with a threshold at 17–18 °C (62.6–64.4 °F) and peak activity at 23–24 °C (73.4–75.2 °F). The proportion of females i...

A holistic approach to developing new corn germplasm that confers multiple insect resistance in various plant tissues at different growth stages was examined. Eight corn germplasm lines were examined for their foliar resistance to fall armyworm [Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae)] and natural enemy attraction at V6–V8 (or 6–8 leaf) stages in 2008 and 2009. Four corn germplasm lines with known levels of resistance to root- and ear-feeding insects [‘CRW3(S1)C6’, ‘B37*H84’, ‘SIM6’ and ‘EPM6’], and four germplasm entries with different levels of S. frugiperda resistance (‘Mp708’, ‘Ab24E’, ‘FAW7061’ and ‘FAW7111’) were evaluated in the study. All plants were manually infested with 15–20 neonate S. frugiperda larvae per plant, and injury was rated 7 and 14 d after infestation. Based on cluster analysis of S. frugiperda injury rating and predator survey data, ‘Mp708’ and ‘FAW7061’ were the most resistant, whereas ‘Ab24E’ and ‘EPM6’ were the most susceptible to fall armyworm fee...

The exotic phloem-feeding emerald ash borer (EAB), Agrilus planipennis, has killed tens of millions of North American ash trees (Fraxinus) since its first detection in the U.S.A. in 2002. Ash trees are killed by larval feeding in the cambial region, which disrupts translocation of photosynthates and nutrients. We observed that EAB larvae feed predominantly downwards in naturally grown green ash trees, a behaviour confirmed in greenhouse-grown black ash seedlings. Furthermore, biomass of larvae feeding downwards was greater than that for larvae feeding upwards. We sought to determine the relative importance of four selection forces (i.e. gravity, moisture content, plant defence, and nutrition) in driving this downward feeding behaviour in this study. The gravity and plant defence (i.e. polyphenols) hypotheses were ruled out because even when seedlings were grown upside down, more EAB larvae moved upwards (towards the root area), and phloem tissue below the feeding site contained higher concentrations of defensive compounds than that above the feeding site.

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.

Tritrophic interactions (plant—herbivore—natural enemy) are basic components of nearly all ecosystems, and are often heavily shaped by bottom-up forces. Numerous factors influence plants’ growth, defense, reproduction, and survival. One critical factor in plant life histories and subsequent trophic levels is nitrogen (N). Because of its importance to plant productivity, N is one of the most frequently used anthropogenic fertilizers in agricultural production and can exert a variety of bottom-up effects and potentially significantly alter tritrophic interactions through various mechanisms. In this paper, the potential effects of N on tritrophic interactions are reviewed. First, in plant-herbivore interactions, N availability can alter quality of the plant (from the herbivore’s nutritional perspective) as food by various means. Second, nitrogen effects can extend directly to natural enemies through herbivores by changes in herbivore quality vis-a-vis the natural enemy, and may even provide herbivores with a defense against natural enemies. Nitrogen also may affect the plant’s indirect defenses, namely the efficacy of natural enemies that kill herbivores attacking the plant. The effects may be expressed via (1) quantitatively and/or qualitatively changing herbivore-induced plant volatiles or other plant features that are crucial for foraging and attack success of natural enemies, (2) modifying plant architecture that might affect natural enemy function, and (3) altering the quality of plant-associated food and shelter for natural enemies. These effects, and their interactive top–down and bottom-up influences, have received limited attention to date, but are of growing significance with the need for expanding global food production (with accompanying use of fertilizer amendments), the widening risks of fertilizer pollution, and the continued increase in atmospheric CO2.

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.

Reducing fertiliser applications can reduce production costs for cotton (Gossypium hirsutum L.) growers, as well as nitrogen (N) leaching into the soil and contamination of surface and ground water. But altered N fertilisation may also affect pests and their natural enemies. In this study, plots with four different levels of fertiliser input (0, 45, 90 and 135 kg ha−1 N) were used to investigate the influence of N on cotton pest and beneficial arthropod populations, and on cotton yield in Tifton, GA, USA. We predicted that (1) N fertilisation will correlate positively with cotton plant growth; (2) increased N fertilisation will increase pest populations because plants with more N will be more nutritious for and attractive to herbivores; (3) populations of beneficial arthropods and predation of pests will decline with increased N fertilisation because of reduced plant signaling; (4) increased N fertilisation will increase pest mortality due to parasitoids because of increased host quality. Cotton plant growth was enhanced by N fertilisation but yield was unaffected. N fertilisation significantly affected some pest arthropods but inconsistently. Mirids were most abundant in the high N treatment in 1 year of the study and cotton aphids were most abundant in the highest N treatment in the other year of the study. Arthropod predators were generally more abundant in the high N treatment but only spiders and Geocoris spp. were significantly affected by N treatment, with highest numbers present in the highest N treatment but the significant differences were each only in a single year. The greatest mortality of sentinel pest eggs (Spodoptera exigua) due to predation occurred under low N conditions. N fertilisation had no significant effects on parasitism of feral or sentinel caterpillars.

Sporadic starvation of herbivorous insect pests, such as the beet armyworm Spodoptera exigua (Hübner), often occurs due to host plant scarcity and dislodgement from host plants resulting from inclement weather, enemy attack, and other perturbations. We hypothesized that (1) starvation of beet armyworm in earlier instars would increase the possibility of undergoing supernumerary instars greater than would be the case for later instars; and (2) starvation of beet armyworm in earlier instars would prolong developmental time more than starvation in later instars. We tested these hypotheses by starving beet armyworm larvae for 48 h in the first, third, and fifth instars, and monitoring their development. Beet armyworms normally have 5 instars before pupation. Significantly more larvae underwent 6 larval instars (20.22 ± 4.35%) when starved for 2 d in the first instar compared with those starved for 2 d in the third (2.0 ± 2.00) and fifth instars (0.0), and relative to unstarved larvae (0.0). The number of instars that S. exigua larvae will experience can be predicted as early as the third instar-head capsule widths in this instar clearly differentiate for those destined to experience 5 and 6 instars. Developmental times of starved larvae also were prolonged compared with unstarved larvae. Developmental times (mean ± SE in days) of larvae starved in the first instar (20.32 ±0.16) were significantly longer than those starved in the third (17.04 ± 0.20) or fifth instars (16.92 ± 0.10), or than unstarved larvae (14.62 ± 0.25). Timing of starvation significantly affected the pupal weight of larvae undergoing 5 instars. This suggests that (1) starvation of 2 d in very early instars (the first instar) increases the pupal weight at the cost of prolonged developmental time; and (2) starvation of 2 d in later instars (the fifth instar) leads to lower pupal biomass, presumably because of inability to compensate for the loss.

Nitrogen (N) is one of the most critical chemical elements for plant and animal growth, exerting a variety of bottom-up effects. Development and oviposition of the beet armyworm, Spodoptera exigua (Hubner) (Lepidoptera: Noctuidae), were studied in relation to varying N fertilization levels (42, 112, 196, and 280 p.p.m.) in cotton [ Gossypium hirsutum L. (Malvaceae)]. Low N fertilization of cotton plants led to reduced plant biomass and a lower percentage of N in leaf blades and in leaf petioles. Development of S. exigua larvae fed with plants with reduced N applications (42 and 112 p.p.m.) was prolonged relative to treatments receiving higher N fertilization. Almost all larvae reared on artificial diets underwent only five instars before pupation. However, most larvae reared on cotton plants, irrespective of N levels, experienced a supernumerary sixth larval instar. Furthermore, significantly more larvae reared on lower N cotton plants underwent supernumerary development compared to larvae reared on higher N cotton plants. Life-time feeding damage per larva ranged from 55 to 65 cm 2 , depending on the nutritional quality of the food plant, although the differences were not statistically significant. Larvae distinguished between cotton plants with various nutritional qualities and fed preferentially on higher N plants. Female moth oviposition choice was also affected by host plant nutritional quality: cotton plants with higher N levels were preferentially chosen by S. exigua females for oviposition. The mechanisms of these effects are unclear, but they can have important implications for population dynamics and pest status of beet armyworms in the field.

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.

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, β-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.

Calosota elongata Gibson (Hymenoptera: Eupelmidae) is a gregarious, ectoparasitic larval parasitoid that was described recently (Gibson 2010) in association with the goldspotted oak borer, Agrilus coxalis Waterhouse [now considered to be Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae)] in its native range in southeastern Arizona. We have discovered this parasitoid outside of its presumptive native range on 1 December 2010 near the Pine Creek Trailhead, Cleveland National Forest in San Diego Co., California, beneath the bark of coast live oak, Quercus agrifolia Nee (Fagaceae), in association with A. auroguttatus.

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).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.A. auroguttatus preferentially attacked previously infested and larger (diameter at breast height15-30 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.

Instar determination of field-collected insect larvae has generally been based on the analysis of head capsule width frequency distributions or bivariate plotting, but few studies have tested the validity of such methods. We used head capsules from exuviae of known instars of the beet armyworm, Spodoptera exigua (Hubner) (Lepidoptera: Noctuidae), to determine the larval instars with the frequency distribution method and Dyar's rule. Head capsule widths of S. exigua ranged from 0.313 to 1.446 mm. The number of instars from the analytical method matched that from the observed data. Based on misclassification rules derived from nonlinear least square fitting of the head capsule width data, the theoretical misclassification rates ranged from 0.62 to 1.92%. Comparing the theoretical distribution to the observed data, the observational misclassification probabilities ranged from 1.18 to 3.03%. There were also 10 head capsules, eight third instars and two fourth instars, not classified into any of the known instars based on the theoretical distributions. Dyar's growth ratios of successive instars ranged from 1.41 to 1.65, and those based on the observed data and theoretical distribution were similar. Both approaches yielded a linear relationship between the natural logarithm of the mean head capsule width and the instar number, which indicates full representation of the larval instars. The results demonstrated that the frequency distribution-based method was robust, although we recommend caution when using such methods to classify head capsules into specific instar classes. Application of computer algorithms should also be accompanied by visual inspection to determine instars from the frequency distribution.

Instead of being passively attacked by insect herbivores, plants possess a myriad of defense mechanisms to protect themselves. These mechanisms function broadly either by directly reducing herbivore fitness (direct plant defense), or by indirectly attracting natural enemies of the herbivores (indirect plant defense). Many biotic and abiotic factors including nitrogen affect the expression of plant defenses. Nitrogen fertilization is by and large the most common agronomic practice in crop production. Anthropogenic use of nitrogen contributes to meeting the food supply for the rapidly increasing in human population. On the other hand, it alters plant defense capability and the global nitrogen cycle, which is increasingly threatening plant and animal biodiversity in managed and natural ecosystems. In this chapter, we first briefly review types of plant defense and ecological and environmental consequences associated with anthropogenic addition of nitrogen into ecosystems. We then review the influences of nitrogen fertilization on plant direct and indirect defenses. Finally, we discuss the possibility of developing new sustainable pest management tactics through nitrogen management minimizing the negative and maximizing the positive components of tritrophic interactions through optimized nitrogen applications in agricultural production.

The biotechnological revolution has forever changed agricultural research and crop production worldwide. Commercial agriculture now includes plants that produce enhanced yield and quality, survival in hostile environmental conditions, manufacture and express defensive toxins, and yield grains with greatly enhanced nutrient fortification. Unprecedented opportunities and ethical challenges created by this new technology are transforming the field of plant protection may be compared with the dramatic changes of an insect undergoes during metamorphosis. In particular, the technology has brought enormous exciting opportunities as well as great challenges that are transforming the research fields of innate (or cisgenic) insect resistance and insect-host plant interactions. Here, we provide a review of the historic milestones in both cisgenic (insertion of genes from sexually compatible stock) and transgenic (insertion of genes from sexually incompatible stock) crop production in agriculture. Biological significance of transgenic technology and the critical role of basic research in crop and human protection are discussed. In addition, three research frontiers that utilize basic and applied research approaches and synergistically combine cisgenes with transgenic technology for broad crop protection (including both yield and quality improvements) are also discussed.

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)-β-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.

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)-β-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)-β-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, β-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.

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.

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.

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.

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.

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.