Stephen R. Clarke

Review of the Operational IPM Program for the Southern Pine Beetle

The operational Integrated Pest Management program for the southern pine beetle (SPB), Dendroctonus frontalis, currently consists of five components: prevention, prediction, detection, evaluation, and direct control. Full implementation of the program is hampered by economic, regulatory, and management constraints, and devastating SPB outbreaks still occur periodically. Recommendations for improving implementation include gaining public acceptance of the program, devising new management strategies, developing area-wide suppression techniques, coordinating efforts among landowners and government agencies, and revising the environmental analysis process. Severe outbreaks will continue without the creation of less susceptible forest conditions and more stable timber markets, plus the development of improved prediction, detection, communication, and suppression practices.

Forest composition following overstory mortality from southern pine beetle and associated treatmentsExperiment Station Project 06-08-109 of the Kentucky Agricultural Experiment Station, Lexington, Kentucky.

Bark beetle caused mortality continues to play a critical role in determining the composition and structure of forests in North America, and revegetation dynamics following these disturbances are poorly understood. We assessed forest composition following southern pine beetle, Dendroctonus frontalis Zimmermann, mortality and associated cut and leave suppression, and compared them with undisturbed loblolly pine, Pinus taeda L., stands in Texas. Abundant hardwood regeneration dominated the understory, with little loblolly pine regeneration following either mortality event. Disturbances eliminated loblolly pine dominance in these even-aged stands, thus stratifying forest structure and apparently increasing stem density (stems·ha–1), richness (species·ha–1), and diversity (ha) in the lower strata. Aspect and elevation, presumably through influences on site moisture, were the primary gradients associated with vegetation variation in the canonical correspondence analyses for new regeneration. Mortality from Den...

Efficacy of two systemic insecticides injected into loblolly pine for protection against southern pine bark beetles (Coleoptera: Curculionidae).

ABSTRACT We evaluated the efficacy of systemic insecticides emamectin benzoate and fipronil for preventing mortality of individual loblolly pines, Pinus taeda L., as a result of attacks by southern pine bark beetles (Coleoptera: Curculionidae, Scolytinae) for two consecutive years in Mississippi (2005–2006) and Alabama (2006–2007). Trees were injected once in the spring of 2005 (Mississippi) or 2006 (Alabama) and then were baited with species-specific bark beetle lures several weeks later. The southern pine beetle, Dendroctonus frontalis Zimmermann, was the target species but was changed to Ips spp. in Mississippi (but not Alabama) the second year because of few southern pine beetle attacks on baited trees. Single injections of emamectin benzoate were effective in reducing tree mortality caused by bark beetles compared with untreated checks. Although less effective overall, fipronil also significantly reduced tree mortality from southern pine beetle compared with the checks during the second year in Alabama. Tree mortality continued well after the lures had been removed. Evaluations of bolts taken from experimental trees killed in 2006 indicated that emamectin benzoate effectively prevented parent bark beetle gallery construction and that fipronil significantly reduced lengths of galleries constructed by adult beetles, brood development, and emergence, compared with checks. In contrast, neither insecticide treatment prevented the bark beetles from inoculating blue stain fungi, Ophiostoma spp., into treated trees.

Effects of Verbenone Dose and Enantiomer on the Interruption of Response of the Red Turpentine Beetle, Dendroctonus valens LeConte (Coleoptera: Scolytidae), to Its Kariomones

Abstract The effects of different doses and enantiomers of verbenone on the interruption of response of red turpentine beetle, Dendroctonus valens LeConte (Coleoptera: Scolytidae), a very aggressive and destructive exotic pest, were conducted using electoantennogram (EAG) assays and behavioral tests in the laboratory and field. No significant differences were found between antennal responses of D. valens by sex, and only one significant difference occurred between their responses to equivalent doses of the (S)-(−) and (R)-(+) enantiomers.Antennal responses to increased doses of (S)-(−)-verbenone reached a plateau at 0.1 μg, whereas the amplitude of response to increased dosages of (R)-(+)-verbenone continued to rise. In a laboratory bioassay, (S)-(−)-verbenone at a stimulus load of 0.1 μg synergized D. valens response to a kairomone blend; but at higher stimulus loads (10 and 100 μg), it functioned as a repellent. In field trials using funnel traps, both (S)-(−)- and (R)-(+)-verbenone bubblecaps inhibited the response of D. valens to the host kairomone blend. These results indicate that verbenone functions as a multipurpose pheromone for D. valens

Biochemical Evidence that Dendroctonus frontalis Consists of Two Sibling Species in Belize and Chiapas, Mexico

ABSTRACT Dendroctonus frontalis Zimmermann (Coleoptera: Curculionidae: Scolytinae) is a major economic pest of pines in the United States, Mexico, and Central America. We report biochemical investigations relevant to the taxonomic status and semiochemistry of two distinct morphotypes of D. frontalis recently detected in the Central American region. Morphotype A beetles (pre-episternal area of prothorax of both sexes smooth, bulging callus on anterolateral margin of prothorax of females) and morphotype B beetles (pre-episternal area of prothorax of both sexes with fine ridges, reduced female callus) collected in infestations in Chiapas, Mexico differed significantly in their production of 10 behaviorally-active compounds occurring in the genus Dendroctonus, including the major pheromone components for D. frontalis. Notably, host-attacking morphotype B females produced hundreds of nanograms of both endo-brevicomin and frontalin, whereas morphotype A females produced similar amounts of frontalin but subnanogram quantities of endo-brevicomin. Reanalysis of a published D. frontalis trapping study in Chiapas indicated that both morphotypes responded in greatest numbers when frontalin and endo-brevicomin baits were both present. In addition, we quantified 18 different cuticular hydrocarbons (the methyl-branched alkane components) from both morphotypes collected in Belize and Chiapas as well as morphotype A beetles from the southeastern United States, and principal component analysis revealed nonoverlapping clusters associated with either morphotype. This evidence of two distinct, complex phenotypes coexisting in the same sites and host trees supports the hypothesis that the D. frontalis morphotypes represent separate species and consequently indicates that the taxonomy of D. frontalis should be re-evaluated in the Central American region.

Electrophysiological and Behavioral Responses of Dendroctonus valens (Coleoptera: Curculionidae: Scolytinae) to Four Bark Beetle Pheromones

ABSTRACT The red turpentine beetle, Dendroctonus valens LeConte (Coleoptera: Curculionidae: Scolytinae), has caused extensive mortality of Pinus tabuliformis Carrière in north central China. The electrophysiological and behavioral activities of the four bark beetle pheromones, frontalin, exobrevicomin, trans-verbenol, and cis-verbenol, singly or in combination with host-produced kairomones, were tested on red turpentine beetles. Both sexes showed a consistent electrophysiological response to the four test compounds. In Y-tube olfactometer bioassays, walking red turpentine beetles selected the host compound (+)-3-carene over any of the test compounds, but significantly higher numbers chose each tested pheromone over a blank control. The four compounds, tested singly or in combination, were not attractive to red turpentine beetles in field trapping studies in 2006 and 2007 and also did not significantly increase trap catch when combined with (+)-3-carene. Frontalin, alone or in combination with exo-brevicomin and trans-verbenol, significantly reduced the attractiveness of (+)-3-carene in 2006 but not in 2007. The possible roles of the pheromones in host and mate finding and selection are discussed.

Two Species within Dendroctonus frontalis (Coleoptera: Curculionidae): Evidence from Morphological, Karyological, Molecular, and Crossing Studies

ABSTRACT Dendroctonus frontalis Zimmermann is considered one of the most important economic and ecological forest pests in the United States, Mexico, and Central America. Recently, two apparent morphological variants of this species were discovered occurring syntopically in Central America and southern Mexico. Morphotype A beetles lack a series of fine parallel ridges on the episternal area of the prothorax that are present on morphotype B. The goal of the present work was to clarify the taxonomic status of the morphotypes of the D. frontalis species complex. Geometric morphometric analyses of seminal rod and spermatheca shape together with the characterization of 16 attributes of external morphology revealed differences in quantitative and qualitative characters that distinguished adults of the two morphotypes from each other as well as from the closely related species Dendroctonus vitei Wood and Dendroctonus mexicanus Hopkins. Karyotype analysis of morphotype B revealed a chromosomal formula (5AA + Xyp) distinct from that found in morphotype A previously reported for D. frontalis (7AA + Xyp). In the laboratory, forced intermorphotype crosses produced F1 progeny but at lower frequency than intramorphotype pairings, and dissections of spermatheca revealed a lower frequency of insemination at least one type of heterotypic cross. Phylogenetic analysis of the D. frontalis species complex based on 786 bp of the cytochrome oxidase I gene indicated that morphotypes B and A are two independent groups with 98% nodal support within D. frontalis. These data provide compelling evidence that the two syntopic morphotypes represent two distinct sibling species.

Field Response of Dendroctonus frontalis (Coleoptera: Scolytinae) to Synthetic Semiochemicals in Chiapas, Mexico

Abstract Dendroctonus frontalis Zimmermann (Coleoptera: Scolytinae) is the most serious pest of pines (Pinus spp.) in Mexico. Conspecifics are attracted to trees undergoing colonization by the aggregation pheromone frontalin, which is synergized by odors of pine oleoresin released from beetle-damaged host tissue. Synthetic racemic frontalin combined with turpentine has been the operational bait used in traps for monitoring populations of D. frontalis in Mexico as well as the United States. Recently, racemic endo-brevicomin has been reported to be a synergist of the frontalin/turpentine bait and as an important component of the aggregation pheromone for D. frontalis populations in the United States. To determine whether racemic endo-brevicomin also might function as an aggregation synergist for the geographically isolated D. frontalis populations of Central America and Mexico, we performed a field trapping trial in Lagunas de Montebello National Park, Chiapas, Mexico, during July and August 2007. The combination of endo-brevicomin (placed either directly on the trap or 4 m away) plus racemic frontalin and turpentine caught at least 5 times more D. frontalis of both sexes than did turpentine either alone or in combination with either frontalin or endo-brevicomin. The addition of endo-brevicomin to the frontalin/turpentine bait also increased the proportion of females trapped. We conclude that the addition of endo-brevicomin might substantially improve the efficiency of the frontalin/turpentine bait for monitoring of D. frontalis in Central America and Mexico. We discuss factors that reconcile our results with previous studies that reported endo-brevicomin to be an attractant antagonist for populations of D. frontalis in Mexico and Honduras.

Implications of Population Phases on the Integrated Pest Management of the Southern Pine Beetle, Dendroctonus frontalis

The southern pine beetle, Dendroctonus frontalis Zimmermann, has three population phases. In the latent phase, southern pine beetle-initiated infestations are absent, and southern pine beetle functions as a secondary bark beetle when present. The outbreak phase results when one or more multi-tree infestations are detected per 1,000 acres of susceptible host type, and the southern pine beetle acts as an aggressive primary colonizer. The intermediate phase consists of population levels between latent and outbreak. The characteristics of these three phases have management implications, and the integrated pest management (IPM) program for the southern pine beetle should be tailored for each phase. Recommendations for each phase are provided and discussed. Prevention and restoration are primary management concerns in the latent phase, whereas suppression takes precedence during an outbreak.

Effect of associated fungi on the immunocompetence of red turpentine beetle larvae, Dendroctonus valens (Coleoptera: Curculionidae: Scolytinae)

Abstract  Dendroctonus–fungus symbioses are often considered as the ideal model systems to study the development and maintenance of ectosymbioses, and diverse interactions, including antagonism, commensalism and mutualism, have been documented between these organisms. The red turpentine beetle, Dendroctonus valens LeConte (Coleoptera: Curculionidae: Scolytinae) is a pine‐killing invasive beetle in northern China. Fungi species Ophiostoma minus, Leptographium sinoprocerum, L. terebrantis and L. procerum were associated with this bark beetle. Antagonistic interactions between D. valens and its associated fungi, such as O. minus and L. sinoprocerum, have been demonstrated, but the underlying causes of this phenomenon are unknown. Here, we first found the two tested fungi species retarded the net weight gain of D. valens larvae after completing 3‐day feeding on their media. Furthermore, we provide direct evidence indicating the effect of associated fungi on the immunocompetence of D. valens larvae to explain the documented antagonism. Our results showed that the activity of phenoloxidase and total phenoloxidase in D. valens larvae were significantly upregulated by two strains of associated fungi, O. minus and L. sinoprocerum as compared with the controls. The phenoloxidase ratio increased significantly in the larvae which had fed for 3 days on media inoculated with O. minus. Because insect immune defenses are costly to be deployed, these results could be explored as one of the underlying mechanisms of the documented antagonism.