Nutritional and Defensive Chemistry of Three North American Ash Species: Possible Roles in Host Performance and Preference by Emerald Ash Borer Adults

Black ash (Fraxinus nigra), green ash (F. pennsylvanica), and white ash (F. americana) are the three most abundant ash species in the northeastern USA. We compared emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), adult performance and preference among seedlings of the three ash species, and then related performance and preference to foli- age nutritional quality and defensive compounds. Longevity of EAB adults reared on green and white ash was found to be greater than on black ash. EAB adult females also seemed to show feeding preference among the three species of ash trees because the total foliage area consumption was greater on green ash and white ash compared to black ash in dual-choice tests; however, the total mass of foliage consumed did not differ. The foliage of all ash species was high in nitrogen and in most macro- and micro-nutrients studied. The patterns of EAB performance and preference did not correspond to any of the individual chemical compounds tested (nitrogen, proteins, most macro- and micro-nutrients, or putative defensive compounds of ash seedlings). Never- theless, greater longevity of EAB adults on green and white ash compared to black ash was probably related to unbalanced nutrients (total nitrogen/total non-structural carbohydrate ratio) of black ash. Putative defensive compounds (i.e., phenolics and protease inhibitors) did not contribute to EAB longevity in this study, probably because (1) EAB adults were able to excrete most of these compounds and (2) their effects were alleviated by high nitrogen levels. More research is needed to elucidate the interactions of nitrogen and carbohydrate levels, and the interactions of nutrient balance and defensive plant allelochemicals on EAB performance and preference

Effects of Visual Silhouette, Leaf Size and Host Species on Feeding Preference by Adult Emerald Ash Borer, \u3ci\u3eAgrilus Planipennis\u3c/i\u3e Fairmaire (Coleoptera: Buprestidae)

The emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) is an invasive species recently established in North America. In large arena bioassays, when given a choice among live green ash, Fraxinus pennsylvanica Marsh and artificial ash saplings that were hidden or exposed from view, beetles preferred live trees (either visible or hidden) compared to artificial trees that had similar visual silhouettes, confirming that olfactory cues are used to locate hosts. Examination of the effect of leaf size revealed that large leaves attracted more beetles than medium-sized leaves that in turn attracted more beetles than small leaves of the same age. Beetles also consumed more of the large leaves in terms of total leaf area than either medium or small leaves, but the proportion of foliage that beetles consumed relative to total available leaf area, did not differ. When newly emerged adults were fed on green and Manchurian ash, Fraxinus mandshurica Rupr., foliage in a no- choice assay, beetles that were given green ash consumed significantly more foliage compared to those that fed on Manchurian ash, but neither longevity nor beetle body weight differed. Our results suggest that while beetles might use olfactory cues to identify suitable hosts, visual cues also play a role in landing and feeding behavior. Manchurian ash might have greater nutritive value or resistance than green ash, necessitating lower consumption and therefore less damage in nature

\u3ci\u3eTomicus Piniperda\u3c/i\u3e (Coleoptera: Scolytidae): Is Shoot Feeding Required for Reproductive Maturation?

The pine shoot beetle, Tomicus piniperda (Coleoptera: Scolytidae), is a univoltine pest of pine in its native range of Europe and Asia. Tomicus piniperda is now widely established in the Great Lakes region and poses a potentially significant threat to other pine-producing areas in North America. An unusual aspect of the life history of T. piniperda is the extended period of maturation-feeding that takes place in the shoots of living pine trees and subsequent overwintering before adults reproduce the following year. We investigated the extent to which shoot-feeding is required by newly-emerged T. piniperda before introduction into Scotch pine (Pinus sylvestris) logs and before any overwintering, in order for successful reproduction to occur. Tomi­cus piniperda F1 adults successfully reproduced in pine logs in the laboratory after either no shoot-feeding or after 2 to 10 weeks of shoot-feeding. Thus, it is theoretically possible for T. piniperda to be multivoltine, yet it remains univoltine

Evaluation of Three Trap Types and Five Lures for Monitoring \u3ci\u3eHylurgus Ligniperda\u3c/i\u3e (Coleoptera: Scolytidae) and Other Local Scolytids in New York

Hylurgus ligniperda (Coleoptera: Scolytidae) is a pine (Pinus spp.) pest native to Eurasia and northern Africa. In December 2000, an established population of H. ligniperda was discovered in Monroe County, New York. When surveys were initiated to determine the distribution of H. ligniperda, questions arose regarding the most effective trap and lure for survey purposes. We conducted a study in April-May 2001 to compare the effectiveness of commercially available scolytid traps and lures for attracting and capturing H. ligniperda. Traps tested included: 1) 12-unit Lindgren funnel trap, 2) Intercept panel trap, and 3) Theysohn slot-trap. Lures tested included: 1) α-pinene high release (750 mg/day) and ethanol (280 mg/ day), 2) α-pinene low release (300 mg/day) and ethanol, 3) β-pinene high release (2000 mg/day) and ethanol, 4) α-pinene low release, and 5) the “exotic bark beetle lure” [ipsdienol (0.15 mg/day), cis-verbenol (0.35 mg/day), and methylbutenol (10 mg/day)]. All three trap designs captured H. ligniperda, however, the Lindgren funnel trap caught significantly higher numbers. Capture rates of Tomicus piniperda (Coleoptera: Scolytidae) and Hylastes opacus were highest in Lindgren funnel traps; whereas Orthotomicus caelatus collections were highest in Theysohn traps. Capture rates of Ips grandicollis and Xyleborinus saxeseni did not vary significantly among trap types. Behavioral differences among scolytid species such as visual stimuli, flight and landing behavior, and host selection may explain some of these differences. Lures containing α-pinene or β-pinene and ethanol were most attractive to H. ligniperda adults, with ethanol and high-release α-pinene attracting the highest numbers in absolute terms. The exotic bark beetle lure was the least attractive lure to H. ligniperda. Attractiveness of the lures tested varied significantly for other Scolytidae, including Dendroctonus valens, H. opacus, Ips calligraphus, I. grandicollis, I. pini, O. caelatus, T. piniperda, and X. saxeseni. These differences likely were due to variation in lure release rates, host preferences, and/or species-specific pheromone attraction

Nutritional and Defensive Chemistry of Three North American Ash Species: Possible Roles in Host Performance and Preference by Emerald Ash Borer Adults

Black ash (Fraxinus nigra), green ash (F. pennsylvanica), and white ash (F. americana) are the three most abundant ash species in the northeastern USA. We compared emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), adult performance and preference among seedlings of the three ash species, and then related performance and preference to foli- age nutritional quality and defensive compounds. Longevity of EAB adults reared on green and white ash was found to be greater than on black ash. EAB adult females also seemed to show feeding preference among the three species of ash trees because the total foliage area consumption was greater on green ash and white ash compared to black ash in dual-choice tests; however, the total mass of foliage consumed did not differ. The foliage of all ash species was high in nitrogen and in most macro- and micro-nutrients studied. The patterns of EAB performance and preference did not correspond to any of the individual chemical compounds tested (nitrogen, proteins, most macro- and micro-nutrients, or putative defensive compounds of ash seedlings). Never- theless, greater longevity of EAB adults on green and white ash compared to black ash was probably related to unbalanced nutrients (total nitrogen/total non-structural carbohydrate ratio) of black ash. Putative defensive compounds (i.e., phenolics and protease inhibitors) did not contribute to EAB longevity in this study, probably because (1) EAB adults were able to excrete most of these compounds and (2) their effects were alleviated by high nitrogen levels. More research is needed to elucidate the interactions of nitrogen and carbohydrate levels, and the interactions of nutrient balance and defensive plant allelochemicals on EAB performance and preference

Effects of Visual Silhouette, Leaf Size and Host Species on Feeding Preference by Adult Emerald Ash Borer, \u3ci\u3eAgrilus Planipennis\u3c/i\u3e Fairmaire (Coleoptera: Buprestidae)

The emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) is an invasive species recently established in North America. In large arena bioassays, when given a choice among live green ash, Fraxinus pennsylvanica Marsh and artificial ash saplings that were hidden or exposed from view, beetles preferred live trees (either visible or hidden) compared to artificial trees that had similar visual silhouettes, confirming that olfactory cues are used to locate hosts. Examination of the effect of leaf size revealed that large leaves attracted more beetles than medium-sized leaves that in turn attracted more beetles than small leaves of the same age. Beetles also consumed more of the large leaves in terms of total leaf area than either medium or small leaves, but the proportion of foliage that beetles consumed relative to total available leaf area, did not differ. When newly emerged adults were fed on green and Manchurian ash, Fraxinus mandshurica Rupr., foliage in a no- choice assay, beetles that were given green ash consumed significantly more foliage compared to those that fed on Manchurian ash, but neither longevity nor beetle body weight differed. Our results suggest that while beetles might use olfactory cues to identify suitable hosts, visual cues also play a role in landing and feeding behavior. Manchurian ash might have greater nutritive value or resistance than green ash, necessitating lower consumption and therefore less damage in nature

Disruptant Effects of 4-Allylanisole and Verbenone on \u3ci\u3eTomicus Piniperda\u3c/i\u3e (Coleoptera: Scolytidae) Response to Baited Traps and Logs

We assessed the inhibitory effects of the host compound 4-allylanisole (release rates = 1 and 2 mg/d in 1994, and 1 and 10 mg/d in 2001) on the response of the pine shoot beetle, Tomicus piniperda (L.), adults to funnel traps baited with the attractant host compound α-pinene (release rate = 150 mg/d) in two pine Christmas tree plantations in Michigan in spring 1994 and two other plantations in spring 2001. In three of the four plantations, all doses of 4- allylanisole significantly reduced T. piniperda attraction to α-pinene-baited traps by 46 to 76%. We also tested the inhibitory effect of the antiaggregation pheromone verbenone (release rates = 2 and 4 mg/d) on T. piniperda attack density on pine bolts (average bolt length was 62 cm and diameter was 19 cm) at three sites (two pine forest stands and one Christmas tree plantation) in Michigan in 1994. Verbenone significantly reduced T. piniperda attack density by 37 to 60% at the two pine stands, but not at the Christmas tree plantation

Behaviors of Adult \u3ci\u3eAgrilus Planipennis\u3c/i\u3e (Coleoptera: Buprestidae)

A 2-year study was conducted in Canada (2003) and the United States (2005) to better understand searching and mating behaviors of adult Agrilus planipennis Fairmaire. In both field and laboratory, adults spent more time resting and walking than feeding or flying. The sex ratio in the field was biased towards males, which tended to hover around trees, likely looking for mates. There was more leaf feeding damage within a tree higher in the canopy than in the lower canopy early in the season, but this difference disappeared over time. In choice experiments, males attempted to mate with individuals of both sexes, but they landed more frequently on females than on males. A series of sexual behaviors was observed in the laboratory, including: exposure of the ovipositor/genitalia, sporadic jumping by males, attempted mating, and mating. Sexual behaviors were absent among 1-3 day-old beetles, but were observed regularly in 10-12 day-old beetles. Females were seen exposing their ovipositor, suggestive of pheromone-calling behavior. No courtship was observed prior to mating. Hovering, searching, and landing behaviors suggest that beetles most likely rely on visual cues during mate finding, although host-plant volatiles and/or pheromones might also be involved

Disruptant Effects of 4-Allylanisole and Verbenone on \u3ci\u3eTomicus Piniperda\u3c/i\u3e (Coleoptera: Scolytidae) Response to Baited Traps and Logs

We assessed the inhibitory effects of the host compound 4-allylanisole (release rates = 1 and 2 mg/d in 1994, and 1 and 10 mg/d in 2001) on the response of the pine shoot beetle, Tomicus piniperda (L.), adults to funnel traps baited with the attractant host compound α-pinene (release rate = 150 mg/d) in two pine Christmas tree plantations in Michigan in spring 1994 and two other plantations in spring 2001. In three of the four plantations, all doses of 4- allylanisole significantly reduced T. piniperda attraction to α-pinene-baited traps by 46 to 76%. We also tested the inhibitory effect of the antiaggregation pheromone verbenone (release rates = 2 and 4 mg/d) on T. piniperda attack density on pine bolts (average bolt length was 62 cm and diameter was 19 cm) at three sites (two pine forest stands and one Christmas tree plantation) in Michigan in 1994. Verbenone significantly reduced T. piniperda attack density by 37 to 60% at the two pine stands, but not at the Christmas tree plantation