Sex pheromone of Nearctic Agriotes mancus and its similarity to that of three Palearctic Agriotes invasive in North America

1. The wheat wireworm, Agriotes mancus (Coleoptera: Elateridae), is a predominant elaterid pest species in the Nearctic region, with a life history and morphology similar to those of Agriotes obscurus, Agriotes lineatus and Agriotes sputator, three Palearctic pest elaterids invasive in North America. Here, we report the identification and field testing of the sex pheromone of A. mancus.
2. We collected headspace volatiles from female beetles on Porapak Q, and analysed aliquots of Porapak extract by gas chromatography with electroantennographic detection (GC-EAD) and by GC-mass spectrometry. In GC-EAD recordings, two esters—geranyl butanoate and geranyl hexanoate—elicited antennal responses from A. mancus males. In field experiments, trap lures containing both geranyl butanoate and geranyl hexanoate afforded large captures of A. mancus males, which were—on average—approximately 30-fold higher than captures in traps baited with a single ester.
3. Traps baited with geranyl butanoate as a single-component lure captured a significant number of Palearctic A. sputator, indicating the establishment of A. sputator in its invaded Nearctic range.
4. With the A. mancus sex pheromone now known, it can be included in the development of pheromone-based programmes to monitor and manage native and invasive Agriotes pests in North America.

     

Antipredator Behavioral Traits of some <Emphasis Type="Italic">Agriotes</Emphasis> Wireworms (Coleoptera: Elateridae) and their Potential Implications for Species Identification

Many insects show a typical antipredator behavior. The two simplest tactics are immobilization and flight. The antipredator behavior of wireworms, the soil-dwelling polyphagous larvae of click beetles, has not yet been investigated. Birds are known to attack wireworms. Our experimental method (picking up and dropping wireworms) mimics the first attack of a predator, such as a bird. In Europe, some Agriotes species are of particular importance as crop pests. As these species differ considerably, in ecology, damage threshold, and pathogen resistance, knowing more about the species specific biology and behavior will support the establishment of successful control measures. In an extensive microcosm study, we observed the antipredator behavior of wireworms belonging to four different Agriotes species (A. lineatus, A. obscurus, A. sordidus, and A. ustulatus) before and during digging into the soil after a startle-stimulus. We recorded wireworm immobility frequency and duration, and locomotor activity at three temperatures. We also analyzed genetic and morphologic attributes of A. lineatus larvae from two different origins. Following behavior types were found reflecting different escape tactics: (a) distinct tonic immobility (A. lineatus), (b) brief/inconsistent tonic immobility (A. obscurus, A. sordidus), and (c) immediate fleeing/burying (A. ustulatus). Additionally, we found small but significant differences in morphology, behavior, and genetics (PCR results) between A. lineatus larvae originating from the Netherlands and those originating from Germany. The biological information we gathered about each species will further increase the understanding of these insects and improve interpretation of future experimental data. In addition, the described behavioral differences between Agriotes obscurus and A. lineatus wireworms may represent a useful additional criterion in morphological species identification.     

Differential responses of three Agriotes click beetle species to pheromone traps

1 Previous work had suggested that adult male click beetles (Agriotes spp.) show differential responses to species specific pheromone traps. This hypothesis was tested using mark–release–recapture methods to estimate the maximum sampling range and the effective sampling area of traps for three species. 2 Captured beetles of the species Agriotes lineatus, Agriotes obscurus and Agriotes sputator were marked to show the direction of release, the distance of the release point from the trap and the replicate. Analysis of variance showed that there were significant differences in recapture rates between species and release distances. There were no significant differences between release direction and replicates. 3 Calculated linear speeds suggested differences in movement rates in the order: A. lineatus > A. obscurus > A. sputator. There were also substantial differences between the species in the maximum sampling ranges and effective sampling areas of the traps. These placed the species in the same order. 4 The results are used to estimate the minimum cost of mass trapping programmes to prevent males from mating, giving values of €165/ha/year (A. lineatus), €247.5/ha/year (A. obscurus) and €2343/ha/year (A. sputator). 5 Implications for the use of pheromone traps in wireworm pest management are discussed. It is concluded that current U.K. recommendations based on the cumulative total catch of the three species over a sampling season can be improved by considering the spatial relationships between the adult trapping system and larval distribution. The current constraint to this is the general inability to separate wireworms into species.     

Linking climate change and insect pest distribution: an example using Agriotes ustulatus Shall. (Coleoptera: Elateridae)

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Assessment of the range of attraction of pheromone traps to Agriotes lineatus and Agriotes obscurus

• 1 The range of attraction of YATLOR pheromone traps was studied to gain information on the number of traps needed for mass trapping of males of two Agriotes species.
• 2 Male click beetles of the species Agriotes lineatus (L.) and Agriotes obscurus (L.) (25–30 individuals per release point) were marked and released at a distance of 2, 5, 10, 15, 20 and 60 m from a pheromone trap both along and opposite to the known prevailing wind direction. Traps were regularly inspected over approximately 1 month. The percentage of recaptured beetles was calculated and analyzed using analysis of variance. Maximum sampling ranges and effective sampling areas were calculated.
• 3 Averaged over all five trials and distances, approximately 40% of the released beetles (A. lineatus and A. obscurus) were recaptured. The percentage recapture of male adults was significantly affected by release distance, whereas no differences were found for species and release direction.
• 4 Males were recaptured from all release points and the percentage recapture decreased (in part significantly) with increasing distance from 76% (2 m) to 35% (15 m) and 9% (60 m), respectively. Most of the beetles were recaptured within the first 3 days after release, independent of the distance, except 60 m. The effective sampling area for A. lineatus was 1089 m² after 12 days and increased to 1735 m² after 30 days. Corresponding values for A. obscurus were considerably higher: 1518 m² for 12 days and 2633 m² for 30 days.
• 5 We conclude that the range of attraction of the pheromone traps for A. lineatus and A. obscurus is comparatively low, providing high percentage recapture only for release distances up to 10 m. Accordingly, any approach targeted on preventing mating by male mass trapping would require a dense network of pheromone traps.

     

Large scale Agriotes spp. click beetle (Coleoptera: Elateridae) invasion of crop land from field margin reservoirs

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Distribution of two European elaterids,Agriotes obscurus and A. lineatus in British Columbia: New records,and potential implications of their dispersal

Two European click beetle species, Agriotes obscurus L. (AO) and A. lineatus L. (AL) have become serious pests of agriculture in the Fraser Valley and Vancouver Island areas of British Columbia (BC), but prior to the start of this survey (2017) it was unknown if they had established elsewhere in the Province. Pitfall traps baited with sex pheromone of AO and AL were placed throughout BC in 2017–2019 and collected a total of 4,988 AO and 20,103 AL beetles in 150 trap pairs. Both species were found to be distributed throughout southern BC, including high numbers in Creston, Kelowna, Pemberton, and Salmon Arm. Traps placed in northern BC, northern Alberta, northern Idaho, or the Willamette valley in western Oregon did not collect AO or AL. Traps were also deployed for A. sputator L., a related European species that has become a serious pest of potato in eastern Canada, but this species was not collected. AO and AL traps placed in the Pemberton valley of BC also collected 474 A. ferrugineipennis (LeConte), and traps placed in the northern Okanagan valley collected 75 A. oregonensis Becker. Both are native Agriotes species not closely related to AO or AL; this is the first time A. oregonensis was collected in Canada. We discuss the potential implications of the spread of AO and AL to other agricultural areas in BC and beyond.     

The effect of simulated seasonal temperatures on Metarhizium brunneum-associated mortality in Agriotes spp. click beetles,and a degree-day infection model

Entomopathogens tend to have a slow speed of kill when used for targeting agricultural insect pests. Relating temperature as a driver of this speed is important to predict pest mortality, and extending this to a degree-day infection model has rarely been studied. Many species of wireworms (Coleoptera: Elateridae), the larvae of click beetles, are subterranean and generalist agricultural pests that can be difficult to control with pesticides. Targeting adult beetles, however, may be an effective method to reduce larval recruitment. Metarhizium brunneum Petch (Hypocreales), an entomopathogenic fungus, kills click beetles but the mortality rate and speed of kill are expected to vary according to temperature. Using a thermal gradient plate to simulate daily oscillating temperatures in Agassiz, British Columbia, Canada, for April, May, and June, the effectiveness of M. brunneum strains LRC112 and F52 in causing mortality to Agriotes obscurus (L.) and Agriotes lineatus (L.) click beetles was studied in the laboratory. Mortality was fastest in beetles exposed to June temperatures and slowest in those exposed to April temperatures, with differences among beetle species × M. brunneum strain combinations. Higher temperatures resulted in more rapid mycelial outgrowth and conidiation in beetle cadavers, with only A. obscurus infected with M. brunneum LRC112 attaining near 100% conidiation. The number of degree days required to kill 50% of the beetles (LDD₅₀) was least for A. obscurus infected with M. brunneum LRC112 (176) followed by A. obscurus × M. brunneum F52 (212), A. lineatus × M. brunneum LRC112 (215), and A. lineatus × M. brunneum F52 (292). Hypothetical calculations showed that M. brunneum exposure earlier in the season resulted in a longer time to kill 50% of the beetles (LT₅₀) but the earliest LT₅₀ calendar date. Later M. brunneum exposure dates resulted in lower LT₅₀'s, but later LT₅₀ dates. This conceptual work demonstrates that daily temperature oscillations, seasonality, and degree days must be considered to predict the efficacy and speed of kill of different fungal entomopathogen strains when targeting different click beetle species.     

Spatial clustering of Diabrotica virgifera virgifera and Agriotes ustulatus in small-scale maize fields without topographic relief drift

The soil‐living larvae of Diabrotica virgifera virgifera LeConte (Coleoptera: Chrysomelidae) and Agriotes ustulatus Schaller (Coleoptera: Elateridae) can cause economic damage to maize roots, Zea mays L. (Poaceae). This study investigated the spatial clustering of both pests in four small‐scale maize fields in southern Hungary, where clustering had been observed but not expected due to the lack of topographic relief drifts and soil structuring. Between 2000 and 2002, numbers of D. v. virgifera larvae and adults and of A. ustulatus larvae were determined at four randomly chosen georeferenced maize plants in each of 24 plots per field. Soil moisture, soil bulk density, and vegetational characteristics were assessed. Moran's I test for spatial autocorrelations, semivariogram analyses, and interpolated mapping revealed that D. v. virgifera larvae and adults were spatially clustered in 67 and 50% of cases, respectively. Larvae of A. ustulatus were clustered in 75% of cases. Diabrotica virgifera virgifera larval distributions were mainly determined by increasing weed density (negative correlation), in particular with high densities of Cirsium arvense (L.) (Asteraceae), as well as by increasing soil moisture (negative correlation). Adult distributions of D. v. virgifera were mainly determined by the density distribution of flowering maize. They were moreover correlated with larval distribution and with the adult distribution of the previous year. The density distributions of male adults differed from those of females. Female density was additionally correlated with higher soil moisture and Poaceae density, e.g., with Sorghum halepense (L.) Pers. No relation was found between the larvae of A. ustulatus and D. v. virgifera. Agriotes ustulatus larval distributions were mainly determined by vegetational cover (correlation with less cover). Conclusively, male and female D. v. virgifera adults, larvae of D. v. virgifera, and larvae of A. ustulatus will display different spatial clustering even within ostensibly homogeneous habitats of flat small‐scale maize fields.     

Population dynamics of wireworms (Coleoptera,Elateridae) in arable land after abandonment

The community of elaterid larvae of three sites (field and two fallows), representing different stages of secondary succession, were studied using soil sampling from 1986 to 1993. All three sites were abandoned arable land: a field cultivated until 1991, a fallow I abandoned in 1986, and a fallow II abandoned about 1976. The fallow II was used as a meadow after abandonment and was regularly mown until 1985, when cultivation stopped. Six species of Elateridae larvae were found at all three study sites. In the field, Agriotes obscurus, Athous niger, Athous subsuscus, Dalopius marginatus and Athous vittatus were found, A. niger and A. obscurus being the most abundant species. During cultivation, larval densities were very low, however, larval abundance increased up to 8.8 ± 8.3 ind. m^-2 when cultivation stopped. Only small A. obscurus larvae were found during cultivation, whereas larval size increased after abandonment. In fallow I A. obscurus, A. niger, A. subsuscus, D. marginatus, and Agrypnus murinus were found and the average annual abundance fluctuated between 0.8 ± 1.4 to 40.8 ± 10.9 ind. m^-2 with A. obscurus being the most abundant species. Fallow II supported the highest densities of wireworms from all plots studied (71.2±35.2 to 280.0±24.8 ind. m^-2). A. obscurus, A. niger, A. subsuscus and D. marginatus were found in fallow II. The abundance of all larval Elateridae as well as the dominant species A. obscurus decreased during the study period, while simultaneously the occurrence of small sized A. obscurus larvae decreased. The frequency of cultivation and time elapsed since last cultivation appeared to be the most important factors affecting elaterid occurrence in the field and fallow I. In fallow II, the decrease in abundance correlates with the accumulation of a dense litter layer, which may correspond with soil surface structure, plant community changes or predator pressure.     

Considérations systématiques et biologiques sur quelques espèces d'Entomophthora du groupeSphaerosperma pathogènes d'insectes [Phycomycètes:Entomophthoraceae]

Résumé Des souches d'Entomophthora du groupesphaerosperma isolées respectivement de chenilles deTortrix viridana L., d'imagos d'Agriotes sputator L. et de nombreuses espèces d'Aphididae doivent être classées dans trois espèces distinctes:E. sphaerosperma Fres.,E. elateridiphaga Turian etE. phalloides Batko. Les différences essentielles concernent la dimension et la forme des conidies primaires et celles des conidies formées à l'extrémité de tubes capillaires ainsi que les exigences nutritives.

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Summary Entomophthora strains belonging to thesphaerosperma group were isolated from caterpillars ofTortrix viridana L., from adults ofAgriotes sputator L. and from various Aphid species. They respectively belong to 3 separated species ofEntomophthora: E. sphaerosperma Fres. s.str., E. elateridiphaga Turian (originally described as subspecies ofsphaerosperma), E. phalloides Batko. The shape of the secondary conidia which are formed at the top of capillary tubes furnish the best character for discrimination.E. sphaerosperma grows very fast on various media including Sabouraud;E. elateridiphaga grows slowly on Sabouraud andE. phalloides requires adjunction of egg yolk.

     

Mass trapping wild Agriotes obscurus and Agriotes lineatus males with pheromone traps in a permanent grassland population reservoir

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Mark–recapture of Agriotes obscurus and Agriotes lineatus with dense arrays of pheromone traps in an undisturbed grassland population reservoir

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Stability of the microbial population in the gut of omnivorous wireworms (Coleoptera,Elateridae)

The variability of the microbial population structure of the gut of omnivorous wireworms Agriotes obscurus (L) and Selatosomus aeneus (L) was studied. The limits of intra- and interspecific and intersite variation were determined. The stability of the microbial composition of the gut allows us to reveal the list of obligate saprotrophs (with 95% probability) using only five replications. In the case of S. aeneus, the influence of starvation and diet change was studied. Starvation changed the microbial population structure, while the diet did not. The results confirm that omnivorous wireworms have a stable gut microbial population, which suggests an advanced mutualistic relationship between wireworms and their gut bacteria, possibly assisting in digestion and providing for ecological flexibility of wireworms.     

Predation of Thereva nobilitata (Fabricius) (Diptera: Therevidae) on Agriotes obscurus L. (Coleoptera: Elateridae)

Little is known about the natural enemies of wireworms (Coleoptera: Elateridae), but there are frequent anecdotal reports of (usually unnamed) stiletto fly larvae (Diptera: Therevidae) preying on various species. We observed larvae of Thereva nobilitata (Fabricius) feeding on larvae of the dusky wireworm, Agriotes obscurus L., during the summer of 2011, in Agassiz, British Columbia. This finding is of interest as: both the predator and the wireworm are introduced species to this area from Europe; T. nobilitata is uncommon in North America; and this predator has not been associated with any wireworm species previously. We observed that larvae of male and female T. nobilitata will feed on various sizes of A. obscurus larvae, most feeding being carried out by the smallest T. nobilitata larvae. These findings suggest future work should assess the potential for therevid larvae as top‐down regulators of Agriotes larvae under field conditions.     

Laboratory and field evaluation of entomopathogenic nematodes for control of Agriotes obscurus (L.) (Coleoptera: Elateridae)

The susceptibility of the dusky wireworm, Agriotes obscurus (L.) (Coleoptera: Elateridae), to different species and strains of entomopathogenic nematodes was tested in a virulence assay in the laboratory. Larvae were exposed to different nematode doses of 50 and 100 IJs/cm². At a dose of 50 IJs/cm², only a commercial strain Heterorhabditis bacteriophora Poinar and the native strain Steinernema carpocapsae (Weiser) B14 caused increased mortality compared with the control (11.1% and 13.3% mortality, respectively). At the higher dose tested, all strains (except Steinernema sp. D122) were virulent to A. obscurus larvae. Steinernema carpocapsae B14 caused higher mortality of wireworm (75.6%) and was used for the assay conducted in cages, with a dose of 100 IJs/cm², in field conditions. The results showed that S. carpocapsae B14 controlled 48.3% of A. obscurus larvae, demonstrating that some entomopathogenic nematodes have the potential to control larvae of A. obscurus. However, further work is needed to improve their efficacy.     

Female Responses to Synthetic Pheromone and Plant Compounds in <Emphasis Type="Italic">Agriotes brevis</Emphasis> Candeze (Coleoptera: Elateridae)

Traps baited with synthetic pheromone components of Agriotes brevis [geranyl butanoate + (E,E)-farnesyl butanoate] captured significantly higher numbers of not only male, but also female beetles, compared to unbaited controls. Catches of both sexes showed a clear positive relationship with increasing doses. In electroantennogram tests, antennal responses of females and males to a number of known Agriotes pheromone components, identified from pheromone glands, showed a similar trend, with geranyl butanoate eliciting the strongest responses. This suggests that the female and male A. brevis antennae are similar with respect to the perception of pheromone compounds, and female beetles have the sensory capabilities to perceive the pheromone components which they produce. Addition of the plant-derived compounds (Z)-3-hexenyl acetate, methyl benzoate, (Z)-3-hexenol and methyl salicylate (identified earlier from foliage as attractive for A. brevis) to the synthetic pheromone significantly increased catches. All the above results suggest that geranyl butanoate and (E,E)-farnesyl butanoate are constituents of an aggregation pheromone of A. brevis, in contrast to the general view of click beetle pheromones being “classical” sex pheromones. These findings could be useful for more precise monitoring and forecasting of damage, based on female catches.     

中国贵州长腹寄蝇属三新种记述（双翅目：寄蝇科）

记述采自贵州地区的长腹寄蝇属Dolichocoxys Townsend 3个新种:黄基长腹寄蝇D. flavibasis sp. nov.、黑腹长腹寄蝇D. obscurus sp. nov.和短柄长腹寄蝇D. brevis sp. nov.。文中附所有新种的详细描述、鉴别特征图、近缘关系的讨论及中国长腹寄蝇属分种检索表。新种的模式标本保存于贵州省安顺市疾病预防控制中心医学昆虫研究室。     

Carabid populations in field beans and their effect on the population dynamics of Sitona lineatus (L.)

The abundance and diversity of potential carabid predators of Sitona lineatus (L.) in field beans were monitored between 1980 and 1983. The absolute abundance of the large predatory species was determined in mark-recapture experiments and a radio-labelling technique was used to estimate the level of predation in the field. Using previously published population data for S. lineatus, the importance of predation in the population dynamics of the weevil was estimated. The mortality of weevil larvae due to predation varied from 0.6%– 10.5% while that of adult weevils varied from 2.6%– 23.8%. It appeared that carabids played a significant role in the population dynamics of S. lineatus and, in years when abundant, could reduce the population of larvae and overwintering adults by more than 30%.     

Preventive application of an entomopathogenic fungus in cover crops for wireworm control

Efficacy of the Metarhizium brunneum Petch (Hypocreales: Clavicipitaceae) strain ART2825 for control of wireworms (Agriotes obscurus (L.), Coleoptera: Elateridae) was examined in a semi-field pot experiment. Pots were treated in late summer during sowing of spring oat as a cover crop. Survival of wireworms was assessed four weeks after their release in October 2013, and 30 weeks after release in April 2014. Viability and persistence of the fungus was determined by counting colony forming units from substrate samples and microsatellite analyses of recovered Metarhizium isolates. The number of colonies detected in the substrate in October 2013 increased with increasing concentrations of applied conidia, and no significant reduction was observed at the second evaluation date in April 2014. Increasing conidia application rates significantly increased mycosis and reduced wireworm survival, to a level comparable to that of treatment using insecticide-coated oat seeds. The preventive application of M. brunneum conidia to reduce wireworm populations in cover crops, preceding a damage-sensitive crop like potatoes, may be a promising biocontrol strategy.