Influence of temperature on spring flight initiation for southwestern ponderosa pine bark beetles (Coleoptera: Curculionidae, Scolytinae)

Determination of temperature requirements for many economically important insects is a cornerstone of pest management. For bark beetles (Coleoptera: Curculionidae, Scolytinae), this information can facilitate timing of management strategies. Our goals were to determine temperature predictors for flight initiation of three species of Ips bark beetles, five species of Dendroctonus bark beetles, and two genera of bark beetle predators, Enoclerus spp. (Coleoptera: Cleridae) and Temnochila chlorodia (Mannerheim) (Coleoptera: Ostomidae), in ponderosa pine forests of northcentral Arizona. We quantified beetle flight activity using data loggers and pheromone-baited funnel traps at 18 sites over 4 yr. Ambient air temperature was monitored using temperature data loggers located in close proximity to funnel traps. We analyzed degree-day accumulation and differences between minimum, average, and maximum ambient temperature for the week before and week of first beetle capture to calculate flight temperature thresholds. Degree-day accumulation was not a good predictor for initiation of beetle flight. For all species analyzed other than D. adjunctus Blandford, beetles were captured in traps only when springtime temperatures exceeded 15.0 degrees C. D. adjunctus was collected when maximum temperatures reached only 14.5 degrees C. Once initial flights had begun, beetles were often captured when maximum ambient air temperatures were below initial threshold temperatures. Maximum and average air temperatures were a better predictor for beetle flight initiation than minimum temperature. We establish a temperature range for effective monitoring of bark beetles and their predators, and we discuss the implications of our results under climate change scenarios.     

Trap type, chirality of alpha-pinene, and geographic region affect sampling efficiency of root and lower stem insects in pine.

Root and lower stem insects cause significant damage to conifers, vector phytopathogenic fungi, and can predispose trees to bark beetle attacks. The development of effective sampling techniques is an important component in managing these cryptic insects. We tested the effects of trap type and stereochemistry of alpha-pinene, in combination with ethanol, on catches of the root colonizing weevils (Coleoptera: Curculionidae) Hylobius spp. [mostly Hylobius pales (Herbst)], and Pachylobius picivorus (Germar), the root colonizing bark beetle (Coleoptera: Scolytidae) Hylastes porculus Erickson, and the lower stem colonizing bark beetle Dendroctonus valens (LeConte). We tested for inter-regional differences by conducting similar field assays in the northern (Wisconsin) and southern (Louisiana) United States. The more effective trap type varied with region. Root weevils were caught primarily in pitfall traps in Wisconsin, whereas they were caught mostly in lower stem flight traps in Louisiana. In Wisconsin, root colonizing bark beetles were also caught primarily in pitfall traps, but lower stem colonizing bark beetles were caught primarily in lower stem flight traps. The root feeding weevils preferred (-) over (+)-alpha-pinene in both regions. Some exceptions relating to trap type or gender occurred in southern populations. The two root and lower stem colonizing bark beetles in Wisconsin showed no preference between (+) and (-)-alpha-pinene in combination with ethanol. No bark beetles were caught in the south. Our results suggest that modifying trap type and enantiomeric ratios of monoterpenes for different insect groups and in different regions can improve sampling efficiency for these important pests.     

Survey and phylogenetic analysis of culturable microbes in the oral secretions of three bark beetle species

In a recent study, we reported a previously undescribed behavior in which a bark beetle exuded oral secretions containing bacteria that have antifungal properties, and hence defend their galleries against pervasive antagonistic Hyphomycete fungi. Actinobacteria, a group known for their antibiotic properties, were the most effective against fungi that invade the spruce beetle galleries. In the present study, we describe the isolation and identification of microorganisms from oral secretions of three bark beetles (Coleoptera: Curculionidae: Scolytinae): the spruce beetle, Dendroctonus rufipennis Kirby, the mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini Say. Bacteria isolated from these three species span the major bacterial classes α-, β-, and γ-Proteobacteria, Firmicutes, Bacteroidetes, and Actinobacteria, except for D. ponderosae , which yielded no α-proteobacteria or Bacteroidetes isolates. Spruce beetles and pine engraver beetles had similar numbers of α-proteobacteria isolates, but pine engravers yielded twice as many Bacteroidetes isolates as spruce beetles. In contrast, mountain pine beetles yielded more isolates in the β- and γ-proteobacteria than spruce beetles and pine engravers. The highest percentage of Actinobacteria was obtained from spruce beetles, followed by pine engravers and mountain pine beetles. All of the fungal isolates obtained from the three beetle species were Ascomycetes. The greatest fungal diversity was obtained in spruce beetles, which had nine species, followed by pine engravers with five, and mountain pine beetles with one.     

Influence of elevation on bark beetle (Coleoptera: Curculionidae, Scolytinae) community structure and flight periodicity in ponderosa pine forests of Arizona

We examined abundance and flight periodicity of five Ips and six Dendroctonus species (Coleoptera: Curculionidae, Scolytinae) among three different elevation bands in ponderosa pine (Pinus ponderosa Douglas ex. Lawson) forests of northcentral Arizona. Bark beetle populations were monitored at 10 sites in each of three elevation bands (low: 1,600-1,736 m; middle: 2,058-2,230 m; high: 2,505-2,651 m) for 3 yr (2004-2006) using pheromone-baited Lindgren funnel traps. Trap contents were collected weekly from March to December. We also studied temperature differences among the elevation bands and what role this may play in beetle flight behavior. Bark beetles, regardless of species, showed no consistent elevational trend in abundance among the three bands. The higher abundances of Ips lecontei Swaine, I. calligraphus ponderosae Swaine, Dendroctonus frontalis Zimmerman, and D. brevicomis LeConte at low and middle elevations offset the greater abundance of I. knausi Swaine, D. adjunctus Blandford, D. approximatus Dietz, and D. valens LeConte at high elevations. I. pini (Say) and I. latidens LeConte were found in similar numbers across the three bands. Flight periodicity of several species varied among elevation bands. In general, the flight period shortened as elevation increased; flight initiated later and terminated earlier in the year. The timing, number, and magnitude of peaks in flight activity also varied among the elevation bands. These results suggest that abundance and flight seasonality of several bark beetles are related to elevation and the associated temperature differences. The implications of these results are discussed in relation to bark beetle management and population dynamics.     

Predators and competitors of the mountain pine beetle Dendroctonus ponderosae (Coleoptera: Curculionidae) in stands of changing forest composition associated with elevation

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Lack of genetic differentiation in aggressive and secondary bark beetles (Coleoptera: Curculionidae, Scolytinae) from Arizona

Bark beetles (Coleoptera: Curculionidae, Scolytinae) play an important role as disturbance agents in ponderosa pine (Pinus ponderosa Douglas ex Lawson) forests of Arizona. However, from 2001 to 2003, elevated bark beetle activity caused unprecedented levels of ponderosa pine mortality. A better understanding of the population structure of these species will facilitate analysis of their dispersal patterns and improve management strategies. Here, we use fluorescently labeled amplified fragment length polymorphism (fAFLP) analysis to resolve genetic variation among and within sampling locations in northcentral Arizona of Ips pini (Say), Dendroctonus brevicomis LeConte, and D. frontalis Zimmermann. We generated genetic fingerprints for >500 beetle specimens and analyzed genetic diversity. For all species, gene flow estimates among sampling locations were high, and significant population subdivision was not discernible across a large portion of ponderosa pine forests in Arizona. However, a weak relationship was detected with I. pini population structure and elevation. Because of the lack of genetic differentiation detected throughout the large study area, our findings suggest these insects are capable of long distance dispersal and exhibit a high degree of gene flow across a broad region. We conclude that our results are consistent with strong dispersal patterns and large population sizes of all three species.     

Gene genealogies reveal cryptic species and host preferences for the pine fungal pathogen Grosmannia clavigera

Grosmannia clavigera is a fungal pathogen of pine forests in western North America and a symbiotic associate of two sister bark beetles: Dendroctonus ponderosae and D. jeffreyi. This fungus and its beetle associate D. ponderosae are expanding in large epidemics in western North America. Using the fungal genome sequence and gene annotations, we assessed whether fungal isolates from the two beetles inhabiting different species of pine in epidemic regions of western Canada and the USA, as well as in localized populations outside of the current epidemic, represent different genetic lineages. We characterized nucleotide variations in 67 genomic regions and selected 15 for the phylogenetic analysis. Using concordance of gene genealogies and distinct ecological characteristics, we identified two sibling phylogenetic species: Gc and Gs. Where the closely related Pinus ponderosa and P. jeffreyi are infested by localized populations of their respective beetles, Gc is present. In contrast, Gs is an exclusive associate of D. ponderosae mainly present on its primary host‐tree P. contorta; however, in the current epidemic areas, it is also found in other pine species. These results suggest that the host‐tree species and the beetle population dynamics may be important factors associated with the genetic divergence and diversity of fungal partners in the beetle‐tree ecosystems. Gc represents the original G. clavigera holotype, and Gs should be described as a new species.     

Evaluation of funnel traps for characterizing the bark beetle (Coleoptera: Scolytidae) communities in ponderosa pine forests of north-central Arizona

Lindgren funnel traps baited with aggregation pheromones are widely used to monitor and manage populations of economically important bark beetles (Coleoptera: Scolytidae). This study was designed to advance our understanding of how funnel trap catches assess bark beetle communities and relative abundance of individual species. In the second year (2005) of a 3-yr study of the bark beetle community structure in north-central Arizona pine (Pinus spp.) forests, we collected data on stand structure, site conditions, and local bark beetle-induced tree mortality at each trap site. We also collected samples of bark from infested (brood) trees near trap sites to identify and determine the population density of bark beetles that were attacking ponderosa pine, Pinus ponderosa Douglas ex Lawson, in the area surrounding the traps. Multiple regression models indicated that the number of Dendroctonus and Ips beetles captured in 2005 was inversely related to elevation of the trap site, and positively associated with the amount of ponderosa pine in the stand surrounding the site. Traps located closer to brood trees also captured more beetles. The relationship between trap catches and host tree mortality was weak and inconsistent in forest stands surrounding the funnel traps, suggesting that trap catches do not provide a good estimate of local beetle-induced tree mortality. However, pheromone-baited funnel trap data and data from gallery identification in bark samples produced statistically similar relative abundance profiles for the five species of bark beetles that we examined, indicating that funnel trap data provided a good assessment of species presence and relative abundance.     

Global and comparative proteomic profiling of overwintering and developing mountain pine beetle,Dendroctonus ponderosae (Coleoptera: Curculionidae), larvae

BackgroundMountain pine beetles, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae), are native to western North America, but have recently begun to expand their range across the Canadian Rocky Mountains. The requirement for larvae to withstand extremely cold winter temperatures and potentially toxic host secondary metabolites in the midst of their ongoing development makes this a critical period of their lives.ResultsWe have uncovered global protein profiles for overwintering mountain pine beetle larvae. We have also quantitatively compared the proteomes for overwintering larvae sampled during autumn cooling and spring warming using iTRAQ methods. We identified 1507 unique proteins across all samples. In total, 33 proteins exhibited differential expression (FDR < 0.05) when compared between larvae before and after a cold snap in the autumn; and 473 proteins exhibited differential expression in the spring when measured before and after a steady incline in mean daily temperature. Eighteen proteins showed significant changes in both autumn and spring samples.ConclusionsThese first proteomic data for mountain pine beetle larvae show evidence of the involvement of trehalose, 2-deoxyglucose, and antioxidant enzymes in overwintering physiology; confirm and expand upon previous work implicating glycerol in cold tolerance in this insect; and provide new, detailed information on developmental processes in beetles. These results and associated data will be an invaluable resource for future targeted research on cold tolerance mechanisms in the mountain pine beetle and developmental biology in coleopterans.     

Efficacy of systemic insecticides for protection of loblolly pine against southern pine engraver beetles (Coleoptera: Curculionidae: Scolytinae) and wood borers (Coleoptera: Cerambycidae)

We evaluated the efficacy of the systemic insecticides dinotefuran, emamectin benzoate, fipronil, and imidacloprid for preventing attacks and brood production of southern pine engraver beetles (Coleoptera: Curculionidae: Scolytinae) and wood borers (Coleoptera: Cerambycidae) on standing, stressed trees and bolt sections of loblolly pine, Pinus taeda L., in eastern Texas. Emamectin benzoate significantly reduced the colonization success of engraver beetles and associated wood borers in both stressed trees and pine bolt sections. Fipronil was nearly as effective as emamectin benzoate in reducing insect colonization of bolts 3 and 5 mo after injection but only moderately effective 1 mo after injection. Fipronil also significantly reduced bark beetle-caused mortality of stressed trees. Imidacloprid and dinotefuran were ineffective in preventing bark beetle and wood borer colonization of bolts or standing, stressed trees. The injected formulation of emamectin benzoate was found to cause long vertical lesions in the sapwood-phloem interface at each injection point.     

Rapid identification and detection of pine pathogenic fungi associated with mountain pine beetles by padlock probes

Fifteen million hectares of pine forests in western Canada have been attacked by the mountain pine beetle (Dendroctonus ponderosae; MPB), leading to devastating economic losses. Grosmannia clavigera and Leptographium longiclavatum, are two fungi intimately associated with the beetles, and are crucial components of the epidemic. To detect and discriminate these two closely related pathogens, we utilized a method based on ligase-mediated nucleotide discrimination with padlock probe technology, and signal amplification by hyperbranched rolling circle amplification (HRCA). Two padlock probes were designed to target species-specific single nucleotide polymorphisms (SNPs) located at the inter-generic spacer 2 region and large subunit of the rRNA respectively, which allows discrimination between the two species. Thirty-four strains of G. clavigera and twenty-five strains of L. longiclavatum representing a broad geographic origin were tested with this assay. The HRCA results were largely in agreement with the conventional identification based on morphology or DNA-based methods. Both probes can also efficiently distinguish the two MPB-associated fungi from other fungi in the MPB, as well as other related fungi in the order Ophiostomatales. We also tested this diagnostic method for the direct detection of these fungi from the DNA of MPB. A nested PCR approach was used to enrich amplicons for signal detection. The results confirmed the presence of these two fungi in MPB. Thus, the padlock probe assay coupled with HRCA is a rapid, sensitive and reproducible method for the identification and detection of these ophiostomatoid fungi.     

Thinning Jeffrey pine stands to reduce susceptibility to bark beetle infestations in California,U.S.A.

• 1 Bark beetles (Coleoptera: Curculionidae, Scolytinae) are commonly recognized as important tree mortality agents in coniferous forests of the western U.S.A.
• 2 High stand density is consistently associated with bark beetle infestations in western coniferous forests, and therefore thinning has long been advocated as a preventive measure to alleviate or reduce the amount of bark beetle‐caused tree mortality.
• 3 The present study aimed to determine the effectiveness of thinning to reduce stand susceptibility to bark beetle infestations over a 10‐year period in Pinus jeffreyi forests on the Tahoe National Forest, California, U.S.A. Four treatments were replicated three times within 1‐ha square experimental plots. Treatments included thinning from below (i.e. initiating in the smallest diameter classes) to a residual target basal area (cross‐sectional area of trees at 1.37 m in height) of: (i) 18.4 m²/ha (low density thin); (ii) 27.6 m²/ha (medium density thin); (iii) 41.3 m²/ha (high density thin); and (iv) no stand manipulation (untreated control).
• 4 Throughout the present study, 107 trees died as a result of bark beetle attacks. Of these, 71% (75 trees) were Abies concolor killed by Scolytus ventralis; 20.6% (22 trees) were Pinus ponderosa killed by Dendroctonus ponderosae; 4.7% (five trees) were P. jeffreyi killed by Dendroctonus jeffreyi; 1.8% (two trees) were P. jeffreyi killed by Ips pini; 0.9% (one tree) were P. jeffreyi killed by Orthotomicus (= Ips) latidens; 0.9% (one tree) were P. ponderosa killed by both Dendroctonus brevicomis and D. ponderosae; and 0.9% (one tree) were P. jeffreyi killed by unknown causes.
• 5 In the low density thin, no pines were killed by bark beetles during the 10‐year period. Significantly fewer trees (per ha/year) were killed in the low density thin than the high density thin or untreated control. No significant treatment effect was observed for the percentage of trees (per year) killed by bark beetles.

     

Integration of visual and olfactory cues of hosts and non-hosts by three bark beetles (Coleoptera: Scolytidae)

Abstract.  1. There has been a long-standing pre-occupation with how phytophagous insects use olfactory cues to discriminate hosts from non-hosts. Foragers, however, should use whatever cues are accurate and easily assessed, including visual cues.
2. It was hypothesised that three bark beetles, the mountain pine beetle (MPB), Dendroctonus ponderosae Hopkins, the Douglas-fir beetle (DFB), D. pseudotsugae Hopkins, and the western balsam bark beetle (WBBB), Dryocoetes confusus Swaine, integrate visual and olfactory information to avoid non-host angiosperms (e.g. paper birch, trembling aspen), that differ in visual and semiochemical profile from their respective host conifers (lodgepole pine, Douglas-fir, interior fir), and tested this hypothesis in a series of field trapping experiments.
3. All three species avoided attractant-baited, white (non-host simulating) multiple-funnel traps, and preferred attractant-baited black (host-simulating) traps. In experiments combining white, non-host traps with non-host angiosperm volatiles, bark beetles were repelled by these stimuli in an additive or redundant manner, confirming that these species could integrate visual and olfactory information to avoid non-host angiosperms while flying.
4. When antiaggregation pheromones were released from white traps, the DFB and MPB were repelled in an additive-redundant manner, suggesting that beetles can integrate diverse and potentially anomalous stimuli.
5. The MPB demonstrated the most consistent visual preferences, suggesting that it may be more of a 'visual specialist' than the DFB or WBBB, for which visual responses may be more contingent on olfactory inputs.     

Diapause and overwintering of two spruce bark beetle species

Diapause, a strategy to endure unfavourable conditions (e.g. cold winters) is commonly found in ectothermic organisms and is characterized by an arrest of development and reproduction, a reduction of metabolic rate, and an increased resistance to adversity. Diapause, in addition to adaptations for surviving low winter temperatures, significantly influences phenology, voltinism and ultimately population growth. We review the literature on diapause and overwintering behaviour of two bark beetle species that affect spruce‐dominated forests in the northern hemisphere, and describe and compare how these strategies can influence population dynamics. The European spruce bark beetle Ips typographus (L.) (Coleoptera, Curculionidae) is the most important forest pest of Norway spruce in Europe. It enters an adult reproductive diapause that might be either facultative or obligate. Obligate diapausing beetles are considered strictly univoltine, entering this dormancy type regardless of environmental cues. Facultative diapausing individuals enter diapause induced by photoperiod, modified by temperature, thus being potentially multivoltine. The spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) infests all spruce species in its natural range in North America. A facultative prepupal diapause is averted by relatively warm temperatures, resulting in a univoltine life cycle, whereas cool temperatures induce prepupal diapause leading to a semivoltine cycle. An adult obligate diapause in D. rufipennis could limit bi‐ or multivoltinism. We discuss and compare the influence of diapause and overwinter survival on voltinism and population dynamics of these two species in a changing climate and provide an outlook on future research.     

Scramble competition in the southern pine beetle, Dendroctonus frontalis

Abstract .1. The nature of intraspecific competition was investigated in the southern pine beetle, Dendroctonus frontalis , a highly destructive pest of pine forests in the southern U.S.A. Data were analysed from an observational study of naturally-attacked trees, and from field experiments where attack density was manipulated by adding different numbers of beetles to caged trees.
2. The effect of attack density on gallery construction, oviposition, brood survival, and the overall rate of increase was examined, and a flexible model of intraspecific competition used to classify the type of competition (contest or scramble) at different points in the life cycle.
3. The results of these analyses suggest that contest competition occurs during gallery construction and oviposition, in accord with previous work on D. frontalis . Strong scramble competition occurs later on in development, however, and the overall competitive process is better characterized as scramble competition, similar to other bark beetles. Trees with attack densities sufficiently high to produce significant competition are common in the field.