David R. Horton

Further comments on analysis of covariance in insect dietary studies

Raubenheimer & Simpson (1992) recently discussed the advantages of using analysis of covariance in insect dietary studies as an alternative to the more conventional ratio‐based nutritional indices. We expand on some interpretations of Raubenheimer & Simpson and illustrate our points with examples from published and unpublished data sets. Specifically, we show that an ANCOVA on biomass gain incorporating initial biomass as a covariate provides information not immediately available using the analysis suggested by Raubenheimer & Simpson (ANCOVA on final biomass incorporating initial biomass as a covariate). Second, we show that dietary studies in which the covariate (food consumption) is affected by diet can provide information about the relative importance of preingestive effects (e.g., deterrence) and postingestive effects (e.g., antibiosis) on performance. Cautions about this latter use of ANCOVA are discussed.

Tree fruit IPM programs in the western United States: the challenge of enhancing biological control through intensive management.

The seminal work of Stern and his coauthors on integrated control has had a profound and long-lasting effect on the development of IPM programs in western orchard systems. Management systems based solely on pesticides have proven to be unstable, and the success of IPM systems in western orchards has been driven by conservation of natural enemies to control secondary pests, combined with pesticides and mating disruption to suppress the key lepidopteran pests. However, the legislatively mandated changes in pesticide use patterns prompted by the Food Quality Protection Act of 1996 have resulted in an increased instability of pest populations in orchards because of natural enemy destruction. The management system changes have made it necessary to focus efforts on enhancing biological control not only of secondary pests but also of primary lepidopteran pests to help augment new pesticides and mating disruption tactics. The new management programs envisioned will be information extensive as well as time sensitive and will require redesign of educational and outreach programs to be successful. The developing programs will continue to use the core principles of Stern and his co-authors, but go beyond them to incorporate changes in society, technology and information transfer, as needed.

Trophic hierarchies illuminated via amino acid isotopic analysis.

Food web ecologists have long sought to characterize the trophic niches of animals using stable isotopic analysis. However, distilling trophic position from isotopic composition has been difficult, largely because of the variability associated with trophic discrimination factors (inter-trophic isotopic fractionation and routing). We circumvented much of this variability using compound-specific isotopic analysis (CSIA). We examined the 15N signatures of amino acids extracted from organisms reared in pure culture at four discrete trophic levels, across two model communities. We calculated the degree of enrichment at each trophic level and found there was a consistent trophic discrimination factor (~7.6‰). The constancy of the CSIA-derived discrimination factor permitted unprecedented accuracy in the measurement of animal trophic position. Conversely, trophic position estimates generated via bulk-15N analysis significantly underestimated trophic position, particularly among higher-order consumers. We then examined the trophic hierarchy of a free-roaming arthropod community, revealing the highest trophic position (5.07) and longest food chain ever reported using CSIA. High accuracy in trophic position estimation brings trophic function into sharper focus, providing greater resolution to the analysis of food webs.

LOCAL DIFFERENCES IN HOST USE BY TWO POPULATIONS OF THE COLORADO POTATO BEETLE

Local differences in performance on three host species (Solanum sarra- choides, S. rostratum, potato (S. tuberosum)) were demonstrated for two nonagricultural populations of Colorado potato beetle. Populations from north-central and northeastern Colorado showed differential larval growth rate, survival, size, and tendency to diapause. Statistically significant host x population interactions indicated that these differences were host-species dependent. Further analysis suggested that the two populations were most successful on the locally abundant and annually predictable host species of their respective community. Population differences in performance were pronounced on one wild host species (S. sarrachoides); performance was similar on the other host species. Extent of host specificity was related to composition of the flora. The population that was exposed to all three host species in the field showed high rates of survival on all hosts; there was no indication that beetles from this population showed major trade-offs in performance among hosts. The second population showed highly reduced survival and growth on the wild host species that is absent from its immediate range. Implications of these results for addressing questions about the beetles original host shift to potato are discussed. Our results, in particular the success of these nonpest populations on potato, suggest that no major phys- iological changes in performance were necessary in the beetle for colonization of potato.

Spider abundance and diversity in apple orchards under three insect pest management programmes in Washington State, U.S.A.

1 Many apple growers in Washington State, U.S.A. use mating disruption (MD) for control of codling moth, Cydia pomonella (Linnaeus). Fewer applications of synthetic, broad‐spectrum insecticides are made in MD orchards than in orchards under conventional (C) management. Spider abundance and diversity in MD, C and certified organic (O) orchards were compared. Spiders inhabiting the trees (arboreal), the understory vegetation, and the ground surface were studied.

Attraction of male pear psylla, Cacopsylla pyricola, to female-infested pear shoots

Post‐diapause winterform pear psylla, Cacopsylla pyricola (Förster) (Homoptera: Psyllidae), exhibit a highly clumped distribution in late winter in pear orchards. The behaviors leading to clumped distributions in this species are unknown, but could include aggregation for mating activities. Choice tests and assays with an olfactometer were done to test whether male psylla of the overwintering morphotype are attracted to pear shoots infested by post‐diapause females and to shoots previously occupied by females. Paired choice tests in small arenas showed that males accumulated on pear shoots currently occupied or previously occupied by females if those shoots were paired with uninfested shoots or shoots previously occupied only by males. Assays with an olfactometer showed that males were attracted to volatile odors from female‐infested or previously infested shoots. The exact source of the attractants (i.e., the female psylla, the pear shoot, or a combination of these sources) remains to be determined.

Effects of mowing frequency on densities of natural enemies in three Pacific Northwest pear orchards

Effects of mowing frequency on ground cover composition and on numbers of predators, parasitoids, and select phytophagous arthropods in the ground cover of three reduced‐insecticide pear orchards were determined. Concurrent samples taken in the tree canopy (with beating trays) and in the herbicide strips on the orchard floor (with pitfall traps) tested whether counts of natural enemies in these two habitats were also affected by mowing regime. A reduction in frequency of mowing from two to three times per month (= control) to once per month or once per growing season led to increased cover of grasses, broadleaf plants, and broadleaf plants in flower. Sweep net samples of natural enemies in the ground cover were dominated numerically by spiders (Araneae), parasitic Hymenoptera, and predatory Heteroptera, with lesser numbers of other taxa (Syrphidae, Neuroptera, Coccinellidae). Predators and parasitoids showed substantial increases in numbers associated with decreased mowing frequency. Sweep net counts of aphids, Lygus spp. (Heteroptera: Miridae), and leafhoppers/planthoppers, all potential prey of predators, also increased significantly with decreased mowing frequency. In the pitfall samples, only the European earwig (Forficula auricularia L.) (Dermaptera: Forficulidae) exhibited a change in counts associated with mowing treatment; numbers of earwigs in pitfall traps declined as mowing frequency decreased. For the beat tray samples, mean tray counts for most natural enemy taxa were higher in the less frequently mowed plots, but significantly (P < 0.05) so only for two taxa: spiders and a predatory mirid, Deraeocoris brevis (Uhler) (Heteroptera: Miridae). It remains to be determined whether biological control of pests in the tree canopy can be enhanced by manipulating mowing frequency. Questions raised by this study include whether there is extensive movement by natural enemies between the ground cover and tree canopy, and whether plot size affects the likelihood of showing that mowing frequency influences predator densities in the tree canopy.

Experimental Infection of Plants with an Herbivore-Associated Bacterial Endosymbiont Influences Herbivore Host Selection Behavior

Although bacterial endosymbioses are common among phloeophagous herbivores, little is known regarding the effects of symbionts on herbivore host selection and population dynamics. We tested the hypothesis that plant selection and reproductive performance by a phloem-feeding herbivore (potato psyllid, Bactericera cockerelli) is mediated by infection of plants with a bacterial endosymbiont. We controlled for the effects of herbivory and endosymbiont infection by exposing potato plants (Solanum tuberosum) to psyllids infected with “Candidatus Liberibacter solanacearum” or to uninfected psyllids. We used these treatments as a basis to experimentally test plant volatile emissions, herbivore settling and oviposition preferences, and herbivore population growth. Three important findings emerged: (1) plant volatile profiles differed with respect to both herbivory and herbivory plus endosymbiont infection when compared to undamaged control plants; (2) herbivores initially settled on plants exposed to endosymbiont-infected psyllids but later defected and oviposited primarily on plants exposed only to uninfected psyllids; and (3) plant infection status had little effect on herbivore reproduction, though plant flowering was associated with a 39% reduction in herbivore density on average. Our experiments support the hypothesis that plant infection with endosymbionts alters plant volatile profiles, and infected plants initially recruited herbivores but later repelled them. Also, our findings suggest that the endosymbiont may not place negative selection pressure on its host herbivore in this system, but plant flowering phenology appears correlated with psyllid population performance.

Comparative efficacy of Beauveria bassiana, Bacillus thuringiensis, and aldicarb for control of Colorado potato beetle in an irrigated desert agroecosystem and their effects on biodiversity

Five weekly applications of Beauveria bassiana (Balsamo) Vuillemin, a genetically engineered isolate of Bacillus thuringiensis Berliner (Raven®), and aldicarb (Temik®) were compared for control of Colorado potato beetle, Leptinotarsa decemlineata (Say) in an irrigated desert cropping system. B. thuringiensis was applied using low and high label rates (1.17 and 7.0 l ha−1). B. bassiana was applied at 5×1013 spores ha−1. Aldicarb (Rhône‐Poulenc), applied at 3.37 kg a.i. ha−1 provided the greatest beetle control and potato yields (45 metric tons ha−1), but overall lowest biodiversity in nontarget organisms, particularly predatory Heteroptera. Low and high rates of B. thuringiensis produced fair to excellent beetle control, yielded 33 and 40 metric tons ha−1 and enabled good survival in predatory Heteroptera and other nontargets. Plots treated with B. bassiana resulted in poor control of beetles prior to row closure after which fair to good control was achieved. Yield in the Beauveria‐treated plots was 33 metric tons ha−1 and effect on biodiversity was comparable to the Bt‐treated plots. The lowest number of overwintering adult L. decemlineata was found in the plots treated with bacteria and fungi (0.68–0.84 adults/0.03 m−3 of soil) and the highest was found in control and aldicarb plots (3.44 and 1.84 adults/0.03 m3 of soil). Aphids and leafhoppers showed higher densities in plots treated with microbial control agents, but were eliminated from plots treated with aldicarb.

Identification of a Sex Attractant Pheromone for Male Winterform Pear Psylla, Cacopsylla pyricola

Pear psylla, Cacopsylla pyricola (Förster) (Hemiptera: Psyllidae), a major economic pest of pears, uses a female-produced sex attractant pheromone. We compared the chemical profiles obtained from cuticular extracts of diapausing and post-diapause winterform males and females to isolate and identify the pheromone. Post-diapause females produced significantly more of the cuticular hydrocarbon, 13-methylheptacosane, than post-diapause males and diapausing females. In olfactometer assays, conspecific males were attracted to synthetic racemic 13-methylheptacosane, whereas females were not, indicating that the behavioral response to this chemical is sex-specific. Furthermore, 13-methylheptacosane was as attractive to males as a cuticular extract of females, suggesting that this chemical was largely responsible for the female attractiveness. A field study showed that males but not females were attracted to 13-methylheptacosane, confirming the olfactometer results. This study provides evidence that 13-methylheptacosane is a sex attractant pheromone for C. pyricola winterform males. This is the first identification of a sex pheromone in the Psylloidea. Our results open the path to developing monitoring tools and possibly new strategies for integrated pest management of this insect.