Potential of a synthetic aggregation pheromone for integrated pest management of Colorado potato beetle

Abstract

• 1 The relative number of colonizing adult Colorado potato beetles (CPB) Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae) coming to pitfall traps baited with the aggregation pheromone (S)‐3,7‐dimethyl‐2‐oxo‐oct‐6‐ene‐1,3‐diol [(S)‐CPB I] and the use of the pheromone in a trap crop pest management strategy were evaluated in the field for the first time.
• 2 More than five‐fold more adult L. decemlineata were caught in pitfall traps baited with the pheromone compared with controls. However, attraction to the pheromone diminished after 5 days in the field.
• 3 In the trap crop management strategy, more colonizing adults were present in pheromone‐treated rows of potatoes compared with untreated middle rows.
• 4 Significantly fewer L. decemlineata egg masses and larvae were found in potato plots that were bordered by pheromone‐treated rows, or bordered by imidacloprid + pheromone‐treated rows, or rows treated at‐planting with imidacloprid compared with untreated (control) potato plots.
• 5 Densities of L. decemlineata egg masses and larvae and percentage defoliation were significantly lower, and marketable tuber yield significantly higher, in conventional imidacloprid‐treated potatoes compared with all other treatments.
• 6 Although our results demonstrate the potential for use of the aggregation pheromone in the management of L. decemlineata in the field, more research is needed to optimize the release rates of the attractant and incorporate control methods for cohabiting pests.

     

Weak population genetic structure in Eurasian spruce bark beetle over large regional scales in Sweden

The Eurasian spruce bark beetle, Ips typographus, is a major pest, capable of killing spruce forests during large population outbreaks. Recorded dispersal distances of individual beetles are typically within hundreds of meters or a few kilometers. However, the connectivity between populations at larger distances and longer time spans and how this is affected by the habitat is less studied, despite its importance for understanding at which distances local outbreaks may spread. Previous population genetic studies in I. typographus typically used low resolution markers. Here, we use genome‐wide data to assess population structure and connectivity of I. typographus in Sweden. We used 152 individuals from 19 population samples, distributed over 830 km from Strömsund (63° 46′ 8″ N) in the north to Nyteboda (56° 8′ 50″ N) in the south, to capture processes at a large regional scale, and a transect sampling design adjacent to a recent outbreak to capture processes at a smaller scale (76 km). Using restriction site‐associated DNA sequencing (RADseq) markers capturing 1409–1997 SNPs throughout the genome, we document a weak genetic structure over the large scale, potentially indicative of high connectivity with extensive gene flow. No differentiation was detected at the smaller scale. We find indications of isolation‐by‐distance both for relative (F _ST) and absolute divergence (Dxy). The two northernmost populations are most differentiated from the remaining populations, and diverge in parallel to the southern populations for a set of outlier loci. In conclusion, the population structure of I. typographus in Sweden is weak, suggesting a high capacity to disperse and establish outbreak populations in new territories.     

Mitochondrial DNA haplotypes indicate two postglacial re‐colonization routes of the spruce bark beetle Ips typographus through northern Europe to Scandinavia

Species in northern Europe re‐colonized the region after the last glacial maximum via several routes, which could have lingering signatures in current intraspecific trait variation. The spruce bark beetle, Ips typographus, occurs across Europe, and biological differences have been found between southern and northern Scandinavian populations. However, the postglacial history of I. typographus in Scandinavia has not been previously studied at a fine geographical scale. Therefore, we collected specimens across northern Europe and analysed the genetic variation in a quite large mitochondrial fragment (698 bp). A high genetic diversity was found in some of the most northern populations, in the Baltic States, Gotland and central Europe. Detected genetic and phylogeographic structures suggest that I. typographus re‐colonized Scandinavia via two pathways, one from the northeast and one from the south. These findings are consistent with the re‐colonization history of its host plant, Picea abies. However, we observed low haplotype and nucleotide diversity in southern Scandinavian populations of I. typographus, indicating that (unlike P. abies) it did not disperse across the Baltic Sea in multiple events. Further, the divergence among Scandinavian populations was shallow, conflicting with a scenario where I. typographus expanded concurrently with its host plant from a ‘cryptic refugium’ in the northwest.     

Reproductive status and flight activity of the overwintered Colorado potato beetle

Mating behavior of post-diapause Colorado potato beetles, Leptinotarsa decemlineata (Say), was observed within an overwintering site, a rotated potato field, newly colonized potato plants, and under laboratory conditions. The influence of spring mating on beetle flight in the presence and in the absence of host plants was investigated using a computer-linked flight mill system. Diapause was terminated simultaneously in male and female beetles, and the first matings were observed as early as within the first 24 h after the beetles emerged from the soil (60–90 DD accumulated). The beetles mated within the overwintering site, the potato field, and the fields rotated out of potatoes. Mating status did not affect flight behavior of overwintered beetles; however, unfed beetles displayed higher flight activity than fed beetles. Most flight activity took place soon after flight muscle regeneration, and then declined sharply by the 5th day after flight initiation. Mating in or near overwintering sites soon after diapause termination might be an important factor in providing gene flow between insecticide-resistant and insecticide-susceptible Colorado potato beetle populations, and should be considered in designing resistance management plans.     

Effects of bark beetle pheromones on the attraction of Monochamus alternatus to pine volatiles

Abstract We evaluated the attraction of Monochamus alternatus Hope (Coleoptera: Cerambycidae), Dryocoetes luteus Blandford and Orthotomicus erosus Wollaston (Coleoptera: Curculionidae: Scolytinae) to multiple-funnel traps baited with the pine volatiles, ethanol and (+)-α-pinene and the bark beetle pheromones, ipsenol and ipsdienol. M. alternatus were attracted to traps baited with ethanol and (+)-α-pinene but not those baited with ipsdienol and ipsenol. Ipsdienol and ipsenol decreased catches of M. alternatus in traps baited with ethanol and (+)-α-pinene. Traps baited with either binary combinations of ethanol and (+)-α-pinene or ipsdienol and ipsenol were attractive to D. luteus and O. erosus. The addition of ipsenol and ipsdienol to traps baited with ethanol and (+)-α-pinene synergized attraction of O. erosus but not D. luteus.     

Fuels for energy metabolism in the Colorado potato beetle, Leptinotarsa decemlineata Say

During flight, the proline concentration in the flight muscles of Leptinotarsa decemlineata decreases sharply while that of alanine shows a proportional increase. In view of the known dynamics of proline turn over the observed decrease would indicate that proline plays the major role as a mobilizable energy source for flight. To a minor extent carbohydrates such as glycogen and glucose support energy metabolism during flight in the Colorado potato beetle.During a recovery period after flight the proline concentration in the fat body increases sharply within minutes, a feature which would indicate that fat body could synthesize proline maybe at the expense of alanine. Comparison of starvation and flight metabolism reveals that the metabolic changes in the two situations are different.     

How varying pest and trap densities affect Tribolium castaneum capture in pheromone traps

The red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), is an important insect pest in food processing facilities. Pheromone trapping is frequently used to monitor red flour beetle populations in structures; however, the optimal trap density and the relationship between trap captures and beetle density is not known. Two experiments were performed concurrently in environmentally controlled 30‐m² walk‐in chambers to determine the relationship between aggregation pheromone trap captures of red flour beetles and beetle and trap number. In one experiment, beetle density was kept constant at 200 individuals per chamber while trap number was varied from 1 to 8, and in the other experiment trap number remained constant at one per chamber while beetle density varied from 20 to 800 individuals. Results indicated that approximately one out of 23 red flour beetles were captured in a trap. Number of beetles captured in traps increased significantly as beetle density increased; however, the proportion of beetles captured remained consistent across beetle densities with a mean of 4.7 ± 0.6% of individuals captured. Trap captures varied significantly with trap placement within experimental chambers, indicating that subtle differences in the trapping environment can influence trap captures. Data suggested that trap densities of 0.07–0.10 m^?2 (2–3 traps per chamber) would maximize trap capture, whereas a trap density of 0.13 m^?2 (four traps per chamber) would maximize the predictive ability of a trapping equation estimating beetle density from trap captures. Results provide information needed to more thoroughly explore how environmental factors might influence red flour beetle trap capture in the absence of changes in beetle density. Further understanding of these relationships will allow for more accurate assessments of absolute beetle density from pheromone trap capture data.     

Genetic relationships of introduced Colorado potato beetle Leptinotarsa decemlineata populations in Xinjiang,China

The Colorado potato beetle (CPB), Leptinotarsa decemlineata (Say), is an infamous invasive species worldwide that aggressively attacks potato and other Solanaceae crops. CPB was first found in China in 1993 and has since spread across 2.77 × 10⁵ km² in Xinjiang Uygur Autonomous Region. To better understand genetic variation and migration patterns, we used seven polymorphic microsatellite loci to elucidate the genetic relationships and gene flow among 10 CPB populations across Xinjiang. (i) Overall low levels of genetic diversity were detected on the entire population in Xinjiang but most of the diversity was retained among populations during invasion. (ii) The mean pairwise F_ST was low (0.071 ± 0.043) among populations. The genetic differentiation was little (pairwise F_ST 0.038 ± 0.016) between the five interior populations (Wusu, Urumqi, Jimsar, Qitai and Mulei) and Tacheng population. The six populations might come from the same genetic group via Bayesian clustering and were closely related on a neighbor‐joining tree. Combining the history data, the five interior populations may have originated from Tacheng. (iii) Gene flow was frequent, especially among the five interior populations. Individuals from the interior populations could be assigned to Tacheng at higher probabilities (means 0.518 ± 0.127) than vice versa (means 0.328 ± 0.074), suggesting that the beetle population has spread from the border to the interior in Xinjiang.     

Survivability and post‐diapause fitness in a scolytid beetle as a function of overwintering developmental stage and the implications for population dynamics

1. It has recently been suggested that expected increases in temperature might lead to an additional generation per season in bark beetles. Thus, populations would grow more rapidly. However, an additional but not fully developed generation could lead to high winter mortality in the pre‐imaginal stages. 2. Winter survivability and post‐diapause fitness as a function of the overwintering developmental stage of the spruce bark beetle Ips typographus L. were studied along an altitudinal cline. Fitness was tested in terms of weight, lipid content and flight capacity. 3. Mortality was significantly lower and emergence per brood system significantly higher for fully developed adults that entered overwintering than for larvae, pupae or callow adults. 4. Post‐diapause fitness in terms of dry weight and flight capacity was significantly higher in individuals that completed development before winter, and lipid contents also showed a trend for being higher in those individuals. 5. In conclusion, in a scenario where effective temperature sums are not adequate for the complete development of an additional generation, models may overestimate population growth by neglecting increased mortality and reduced post‐diapause fitness. The results highlight the importance of considering life‐history traits and indirect effects in addition to abiotic factors such as temperature when modelling population dynamics.     

Physiological and biochemical analysis of overwintering and cold tolerance in two Central European populations of the spruce bark beetle, Ips typographus

Overwintering success is one of the key aspects affecting the development and outbreaks of the spruce bark beetle, Ips typographus (L.) populations. This paper brings detailed analysis of cold tolerance, and its influence on overwintering success, in two Central European populations of I. typographus during two cold seasons. Evidence for a supercooling strategy in overwintering adults is provided. The lower lethal temperature corresponds well to the supercooling point that ranges between −20 and −22 °C during winter months. The supercooled state is stabilized by the absence of internal ice nucleators and by seasonal accumulation of a mixture of sugars and polyols up to the sum concentration of 900 mM. The cryoprotective function of accumulated metabolites is probably based on increasing the osmolality and viscosity of supercooled body fluids and decreasing the relative proportion of water molecules available for lethal formation of ice nuclei. No activity of thermal hysteresis factors (stabilizers of supercooled state) was detected in hemolymph. Lethal times for 50% mortality (Lts50) in the supercooled state at −5, −10 or −15 °C are weeks (autumn, spring) or even months (winter), suggesting relatively little mortality caused by chill injury. Lts50 at −15 °C are significantly shorter in moist (6.9 days) than in dry (>42 days) microenvironment because there is higher probability of external ice nucleation and occurrence of lethal freezing in the moist situation.     

Effects of heat on the dispersal performance of Ips typographus

Under present climate conditions, Ips typographus (L.) is Europe's most critical disturbance agent for mature Norway spruce (Picea abies). With ongoing climate change, the bark beetle will most probably become more prominent as a pest. The aim of this study was to analyse the dispersal performance of I. typographus under various weather conditions, especially hot days with a maximum air temperature above 30°C. In a field study, marked bark beetles were released from breeding logs and could be retraced in traps distributed across the survey area. With daily collections of the trapped beetles, it was possible to analyse the flight activity and the average flight distance of the bark beetles during hot, moderate and cool days. The numbers of daily catches and the average flight distance during the hottest days (air temperature maximum ≥30°C) did not significantly differ from the moderate days (air temperature maximum ≥22°C and <30°C). The numbers of daily catches and the average flight distance during the cool days (air temperature maximum <22°C) were significantly lower than during hot and moderate days. The results give an insight on the dispersal capacity of I. typographus under climate change driven future conditions. Increased air temperatures do not seem to impair the flight performance of I. typographus. A small proportion of cool days during the swarming period even seems to favour dispersal of I. typographus.     

Effect of diet and feeding history on flight of Colorado potato beetle, Leptinotarsa decemlineata

We evaluated the hypothesis that Colorado potato beetle (Leptinotarsa decemlineata Say) (CPB) flight frequency is related to diet, and that it changes with duration of food unavailability or exposure to poor quality food by exposing adult overwintered and summer CPB populations to an acceptable host plant (conventional foliage), a poor host (insect resistant transgenic foliage expressing Bacillus thuringiensis tenebrionis[Btt] Cry3a toxin) and no host. Exposure to poor host and no host treatments (with or without water) decreased mean daily flight frequencies and the overall number of overwintered CPB flying, but increased the mean daily flight frequency and number of summer population CPB that flew. Overwintered CPB did not react to an absence of plants at emergence whereas summer CPB increased mean daily flight frequencies when plants and water were not available. The flight response to insect resistant foliage was similar to that for starvation treatments in both populations indicating that flight may not be triggered by Btt toxins but by starvation brought on by feeding on poor quality food. Flight was observed in all treatments for the duration of the test with two exceptions; overwintered beetles fed insect resistant foliage ceased flying after day 17 and summer beetles starved without water ceased after day 8 of a 29‐day study.     

Synthetic host volatiles increase efficacy of trap cropping for management of Colorado potato beetle, Leptinotarsa decemlineata (Say)

Abstract 1 The attractiveness of pitfall traps baited with a synthetic host volatile attractant to colonizing adult Colorado potato beetle, Leptinotarsa decemlineata (Say) was evaluated in a field setting. 2 Significantly more postdiapause, colonizing adult L. decemlineata were captured in baited than unbaited pitfall traps. 3 The potential for this synthetic kairomone to enhance the efficacy of trap cropping as a management tool was evaluated by comparing conventionally managed plots with like‐sized plots bordered by either attractant‐treated trap crop or untreated trap crop. 4 More postdiapause, colonizing adults, egg masses and small larvae were present in attractant‐treated trap crops than in untreated trap crops. 5 There were no significant differences in egg mass and small larvae densities between plots bordered by attractant‐treated trap crops and conventionally managed plots, but there were significantly fewer large larvae and adult beetles in conventionally managed plots. 6 Plant canopy area of conventionally managed plots was significantly greater than in plots bordered by either type of trap crop. 7 Yields for conventionally managed plots and plots bordered by attractant‐treated trap crops did not differ, and less insecticide (44%) was applied to plots bordered by attractant‐treated trap crops.     

Climate change alters elevational phenology patterns of the European spruce bark beetle (Ips typographus)

The European spruce bark beetle Ips typographus is the most important insect pest in Central European forests. Under climate change, its phenology is presumed to be changing and mass infestations becoming more likely. While several studies have investigated climate effects across a latitudinal gradient, it remains an open question how phenology will change depending on elevation and topology. Knowing how an altered climate is likely to affect bark beetle populations, particularly across diverse topographies and elevations, is essential for adaptive management. We developed a time‐varying distributed delay model to predict the phenology of I. typographus. This approach has the particular advantage of capturing the variability within populations and thus representing its stage structure at any time. The model is applied for three regional climate change scenarios, A1B, A2 and RCP3PD, to the diverse topography of Switzerland, covering a large range of elevations, aspects and slopes. We found a strong negative relationship between voltinism and elevation. Under climate change, the model predicts an increasing number of generations over the whole elevational gradient, which will be more pronounced at low elevations. In contrast, the pre‐shift in spring swarming is expected to be greater at higher elevations. In comparison, the general trend of faster beetle development on steep southern slopes is only of minor importance. Overall, the maximum elevation allowing a complete yearly generation will move upwards. Generally, the predicted increase in number of generations, earlier spring swarming, more aggregated swarming, together with a projected increase in drought and storm events, will result in a higher risk of mass infestations. This will increase the pressure on spruce stands particularly in the lowlands and require intensified management efforts. It calls for adapted long‐term silvicultural strategies to mitigate the loss of ecosystem services such as timber production protection against rockfall and avalanches and carbon storage.     

Changes in emergence phenology,fatty acid composition,and xenobiotic-metabolizing enzyme expression is associated with increased insecticide resistance in the Colorado potato beetle

The Colorado potato beetle (Leptinotarsa decemlineata) is a major agricultural pest of solanaceous crops. An effective management strategy employed by agricultural producers to control this pest species is the use of systemic insecticides. Recent emphasis has been placed on the use of neonicotinoid insecticides. Despite efforts to curb resistance development through integrated pest management approaches, resistance to neonicotinoids in L. decemlineata populations continues to increase. One contributing factor may be alterations in insect fatty acids, which have multiple metabolic functions and are associated with the synthesis of xenobiotic-metabolizing enzymes to mitigate the effects of insecticide exposure. In this study, we analyzed the fatty acid composition of L. decemlineata populations collected from an organic production field and from a commercially managed field to determine if fatty acid composition varied between the two populations. We demonstrate that a population of L. decemlineata that has a history of systemic neonicotinoid exposure (commercially managed) has a different lipid composition and differential expression of known metabolic detoxification mechanisms relative to a population that has not been exposed to neonicotinoids (organically managed). The fatty acid data indicated an upregulation of Δ⁶ desaturase in the commercially managed L. decemlineata population and suggest a role for eicosanoids and associated metabolic enzymes as potential modulators of insecticide resistance. We further observed a pattern of delayed emergence within the commercially managed population compared with the organically managed population. Variations in emergence timing together with specific fatty acid regulation may significantly influence the capacity of L. decemlineata to develop insecticide resistance.     

Incorporating variability in simulations of seasonally forced phenology using integral projection models

Phenology models are becoming increasingly important tools to accurately predict how climate change will impact the life histories of organisms. We propose a class of integral projection phenology models derived from stochastic individual‐based models of insect development and demography. Our derivation, which is based on the rate summation concept, produces integral projection models that capture the effect of phenotypic rate variability on insect phenology, but which are typically more computationally frugal than equivalent individual‐based phenology models. We demonstrate our approach using a temperature‐dependent model of the demography of the mountain pine beetle (Dendroctonus ponderosae Hopkins), an insect that kills mature pine trees. This work illustrates how a wide range of stochastic phenology models can be reformulated as integral projection models. Due to their computational efficiency, these integral projection models are suitable for deployment in large‐scale simulations, such as studies of altered pest distributions under climate change.     

Colonization of potato fields in eastern North Carolina by Colorado potato beetle

Rotated and non-rotated commercial potato fields were sampled intensively to follow Colorado potato beetle,Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), colonization and subsequent oviposition patterns in the spring of 1990 and 1991. Maximum densities of colonizing adults ranged from 0 to 14,891/ha and maximum egg mass densities ranged from 0 to 48,451/ha. Crop rotation generally resulted in lower potato beetle populations. Regardless of crop rotation management practices, colonization of fields planted in potatoes began at field edges and progressed inward in all fields for both years. Management of potato beetles is discussed in light of these findings.     

The role of semiochemicals in tritrophic interactions between the spruce bark beetle Ips typographus, its predators and infested spruce

Abstract:  Semiochemical interactions between the spruce bark beetle Ips typographus , its predators Medetera setiventris , Thanasimus formicarius and Thanasimus femoralis , and the host Norway spruce, Picea abies , were studied in the field. The chemicals S - cis -verbenol, 2-methyl-3-buten-2-ol, ipsdienol, (+)- α -pinene, (−)- α -pinene, (±)- α -pinene, limonene, camphor and their naturally occuring mixtures were used as trap baits in a multiple-choice design that allowed for comparison of their attractivity for the focal species. Medetera was attracted to both the prey aggregation pheromone and its multifunctional component, ipsdienol. On the contrary, both Thanasimus species responded predominantly to ipsdienol and less to the prey aggregation pheromone. In the case of I. typographus , the attractivity of aggregation pheromone seems to be increased by the addition of a mixture of monoterpenic tree volatiles, and by addition of ipsdienol. Bark beetles and predators showed species-specific responses to volatile mixtures representing different stages of tree decay and different stages of bark beetle colony establishment. These responses correlates with the optimal foraging habitat of each species. None of the predator species responded to 2-methyl-3-buten-2-ol, a substantial component of I. typographus pheromonal bouquet, thus it is hypothesized that only substances of monoterpenic origin attract predators.