Development of a standardized protein immunomarking protocol for insect mark–capture dispersal research

A field study was conducted to test the marking efficiency of broadcast spray applications of protein marks on stationary (represented by cadavers) and free‐roaming lady beetles Hippodamia convergens Guérin‐Méneville that were strategically placed in blooming alfalfa plots. The marks tested included three different concentrations of egg albumin from chicken egg white, casein from bovine milk and trypsin inhibitor from soy milk. The cadaver and free‐roaming beetle treatments served to measure the acquisition and retention of each protein treatment regime by direct contact with the spray solution and by residual contact with protein‐marked residue on alfalfa, respectively. In addition, the vertical distribution of marking efficacy was determined by sampling alfalfa plant tissue and beetle cadavers that were located on the upper and lower portion of the plant canopy. The data indicated that the backpack spray apparatus was very effective at uniformly administering the various protein marks, regardless of the concentration, throughout the entire plant canopy. Also, the free‐roaming beetles readily self‐marked by contact exposure to protein‐treated plants. We also identified concentrations of each protein type that will mark about 90% of the resident beetle population. Moreover, if a mark–capture‐type study only requires two unique protein marks, we determined that concentrations of 25% for egg white and 100% for bovine milk could be used to mark 98% of the population. Our results provide a significant step towards standardizing protein immunomarking protocols for insect mark–capture dispersal research. In addition, we identify several areas of research that are needed to further standardize the protein mark–capture procedure.     

Effects of grafting on host plant resistance in ash (Fraxinus spp.) to emerald ash borer (Agrilus planipennis Fairmaire)

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Visually Mediated ‘Paratrooper Copulations’ in the Mating Behavior of <Emphasis Type="Italic">Agrilus planipennis</Emphasis> (Coleoptera: Buprestidae), a Highly Destructive Invasive Pest of North American Ash Trees

The emerald ash borer, Agrilus planipennis, is a serious invasive pest of North American ash (Fraxinus) trees. In captivity, mating is initiated by beetles at least 10 days old, and appears to be based simply on random contact with a member of the opposite sex. In the field, male A. planipennis search the tree during flight, and attempt to copulate with dead beetles of both sexes pinned to leaves, after descending rapidly straight down onto the pinned beetles from a height of from 30 to 100 cm. All evidence suggests that males find potential mates using visual cues. Equal numbers of feral males approach all ‘dummy’ beetles; however, considerably more time is spent attempting copulation with dead females rather than males, suggesting a contact chemical cue. Sticky traps prepared from dead, pinned EAB capture crawling insects as well as male A. planipennis, at a rate similar to that at which small purple sticky traps of similar overall area capture crawling insects and both sexes of feral EAB.     

Marking Triatoma brasiliensis,Triatoma pseudomaculata and Rhodnius nasutus Nymphs with Trace Elements: Element Persistence and Effects of Marking on Insect Mortality

BackgroundField ecologists often rely on mark-release-recapture (MRR) experiments to estimate population dynamics parameters for a given species. In the case of a medically important taxon, i.e., a disease vector, inferences on species survival and dispersal rates are particularly important as they have the potential to provide insights into disease transmission dynamics in endemic areas. Medical entomologists have traditionally used fluorescent dusts to externally mark the cuticle of insects. However, dust marking is usually restricted to the adult life stage because immature insects lose the mark when they molt.Conclusions/SignificancePrevious evidence shows that there is little or no genetic differentiation between populations at the microgeographic level, which often precludes indirect estimations of dispersal capability based on genetic markers. In such situations, MRR studies are more suitable as they measure insect movement directly from one site to another, instead of effective migration (i.e. gene flow). The determination of a reliable and persistent marking method is the first step towards the development of meaningful ecological estimates through the application of MRR methodology. Here, we have identified trace elements that can be used for mark and recapture studies of three triatomine species in Brazil.     

Application of harmonic radar to analyze dispersal behavior of the Japanese pine sawyer beetle,Monochamus alternatus (Coleoptera: Cerambycidae)

Monochamus alternatus (Hope) is a severe wood‐boring pest in coniferous forests and a major vector of pine wilt disease in East Asia. Harmonic radar is a powerful tool for studying the dispersal behavior of insects and it could be applied to control pine wilt disease. In this study, we validated the application of harmonic radar for analyzing the dispersal behavior of M. alternatus beetles in a field environment. We determined the wing capacities of the beetles and the effects of electronic tagging and marking on their movement, flight ability, survivorship, and food consumption in the laboratory to confirm the suitability of this technique. The detection rate and recovery rate for beetles were analyzed separately using radar on caged pine stands and in the field environment. The results showed that the minimum wing capacity of the Japanese pine sawyer was 24.9 mg, which was seven times the weight of the electronic tag (3.5 mg). Marking and tagging the beetles had no significant adverse effects on their movement, flight capacity, food consumption, and survivorship. The detection rate using the radar system and recovery rate based on visual observations of the beetles in caged pines were both 95.6%. However, in the field environment, the detection and recovery rates were only 55.6% and 37.8% after one week, respectively, and 33.3% and 7.8% after two weeks. Harmonic radar is a promising technique for studying the dispersal behavior of the Japanese pine sawyer, but its performance is not satisfactory and major improvements are required for both the radar system and electronic tags.     

Effects of the emerald ash borer invasion on the community composition of arthropods associated with ash tree boles in Maryland,U.S.A.

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An agent-based modeling approach to represent infestation dynamics of the emerald ash borer beetle

Agent-based modeling (ABM) is a bottom-up approach capable of operationalizing complex systems. The approach can be used to reproduce the spatio-temporal patterns in ecological processes such as insect infestation by representing individual dynamics and interactions between “agents” and their environment from which complex behavior emerges. The emerald ash borer (Agrilus planipennis; EAB) is an invasive species native to south-east Asia which has infested and killed millions of ash trees (Fraxinus sp.) across the eastern United States as well as Ontario and Quebec in Canada. Efforts to model the insect's behavior are ongoing, but current models are limited to approaches that do not address the complexity that emerges from the dynamics between individual beetles and their varying spatial environments. The objective of this study is to develop an ABM to represent the interactions of the EAB and the emerging spatio-temporal pattern of the insect spread. The model is implemented on real datasets from the Town of Oakville, Ontario, Canada from 2008 to 2010. Tree inventory and land use data acquired from the Town of Oakville were used to represent the spatial environment of the EAB agents. The EAB interactions are implemented in the model as subroutines, each representing a stage in the EAB life cycle using a temporal resolution of one day. Model verification was performed based on the literature documenting the life cycle processes of the EAB to represent EAB behavior. The model is calibrated using the rate of spread observed in the Town of Oakville from 2008 to 2009 and is validated using datasets delimiting the spatial extent and severity of EAB infestation in 2009. When comparing simulated and observed data, there is a 72% agreement for the locations of the infestation. This indicates that the developed ABM approach offers a model able to capture the complex behavior of EAB where both the spatial extent and severity of infestation are simulated realistically. The model generates insights about the underlying processes governing EAB behavior, highlights areas of uncertainty in modeling the complex spatio-temporal patterns of EAB infestation, and is a useful tool for forest and pest management.     

Overwintering sites of the Japanese carrion beetle <Emphasis Type="Italic">Eusilpha japonica</Emphasis>

Overwintering sites have recently been a focus of research into adaptive insect behavior in areas where winter is severe; however, little is known about the overwintering sites of most insects, including common ground or carrion beetles. Here, we reveal the overwintering site of Japanese carrion beetles Eusilpha japonica (Motschulsky) (Coleoptera: Silphidae). These beetles occupy various locations between spring and autumn; however, they are not observed in most of these places in winter. To investigate where they overwinter, a thorough scan was made of 109 sites. Overwintering beetles were found at 21 of 73 sites near to trees, and no beetles were observed at 36 sites that were not close to trees. To evaluate the influence of trees on the abundance of overwintering beetles, we established a quadrat consisting of 518 subquadrats in a suitable habitat with varying distances to the nearest trees. We caught 118 overwintering beetles in the quadrat. More beetles existed in subquadrats closer to trees, and analyses that considered spatial autocorrelation revealed a negative correlation between beetle frequency and distance to the nearest tree.     

Comparative efficacy of five types of trap for woodborers in the Cerambycidae, Buprestidae and Siricidae

1 Traps of four new designs were tested against the conventionally used multiple‐funnel trap to determine whether trapping of large wood‐boring insects can be improved in western Canada. All four new traps used a large collecting receptacle containing detergent‐laced water, and three presented a prominent visual silhouette above the receptacle. 2 In total, 27 336 large woodborers were captured from 10 June to 30 September in an experiment in the southern interior of British Columbia, and 4737 from 6 June to 27 July in an experiment in northern Alberta. The woodborers captured in the British Columbia experiment were mainly beetles in the families Cerambycidae (79%) and Buprestidae (15%), and woodwasps in the family Siricidae (6%). Most woodborers, e.g. three Monochamus spp. and Xylotrechus longitarsus (the predominant cerambycids), were captured throughout the summer, with peak captures in August. 3 Cross‐vane, pipe and stacked‐bottomless‐flower‐pot traps were generally superior to pan and multiple‐funnel traps for insects in nine taxa, but cross‐vane traps were the most effective overall, trapping 32% of all insects captured. 4 The large number of target insects captured in a relatively small number of traps in the two experiments suggests that employment of an efficacious trap with a large vertical silhouette and a wide, escape‐proof collecting receptacle could make mass trapping of large woodborers in timber processing areas operationally feasible. 5 Because the most effective traps were unstable in the wind, and the detergent‐laced water captured unacceptably high numbers of small mammals, design modifications are necessary. We are currently developing a wind‐firm trap, with a prominent vertical silhouette, a wide collecting surface, and an escape‐proof, but dry collecting receptacle.     

Effects of SmartWater,a fluorescent mark,on the dispersal,behavior, and biocontrol efficacy of Phytoseiulus persimilis

New marking methods for studying small biocontrol agents (especially predatory mites) are needed because many current techniques are expensive, ineffective or not applicable to small organisms. The objective of this study was to determine whether SmartWater, a liquid and permanent fluorescent dye, can be used to mark Phytoseiulus persimilis for experimentation without any deleterious effects on its dispersal, behavior, reproduction, and biocontrol efficacy. Our results show that there were no significant differences in movement, inter-plant dispersal, feeding behavior, survivability, and reproduction between marked P. persimilis and control individuals sprayed with water. We also found that the SmartWater mark lasted for the duration of the mites’ life, indicating strong durability over time. Marking efficacy may be reduced, due to a trade-off between batch marking efficacy and the possibility of drowning study organisms. However, we feel future research could improve liquid marking techniques that would reduce this risk. Overall, this study concludes that SmartWater could be a useful marking tool for predatory mites in both laboratory and field studies.

     

Studying the movement behavior of benthic macroinvertebrates with automated video tracking

Quantifying and understanding movement is critical for a wide range of questions in basic and applied ecology. Movement ecology is also fostered by technological advances that allow automated tracking for a wide range of animal species. However, for aquatic macroinvertebrates, such detailed methods do not yet exist. We developed a video tracking method for two different species of benthic macroinvertebrates, the crawling isopod Asellus aquaticus and the swimming fresh water amphipod Gammarus pulex. We tested the effects of different light sources and marking techniques on their movement behavior to establish the possibilities and limitations of the experimental protocol and to ensure that the basic handling of test specimens would not bias conclusions drawn from movement path analyses. To demonstrate the versatility of our method, we studied the influence of varying population densities on different movement parameters related to resting behavior, directionality, and step lengths. We found that our method allows studying species with different modes of dispersal and under different conditions. For example, we found that gammarids spend more time moving at higher population densities, while asellids rest more under similar conditions. At the same time, in response to higher densities, gammarids mostly decreased average step lengths, whereas asellids did not. Gammarids, however, were also more sensitive to general handling and marking than asellids. Our protocol for marking and video tracking can be easily adopted for other species of aquatic macroinvertebrates or testing conditions, for example, presence or absence of food sources, shelter, or predator cues. Nevertheless, limitations with regard to the marking protocol, material, and a species’ physical build need to be considered and tested before a wider application, particularly for swimming species. Data obtained with this approach can deepen the understanding of population dynamics on larger spatial scales and of the effects of different management strategies on a species’ dispersal potential.     

Dispersal and influences on movement for Anoplophora glabripennis calculated from individual mark-recapture

We conducted an individual mark‐release‐recapture experiment on the beetle, Anoplophora glabripennis Motchulsky (Coleoptera: Cerambycidae). This invasive beetle has been introduced from Asia to Europe and North America and poses a serious threat to several important species of tree. Eradication efforts may benefit from knowledge of dispersal behaviour. Trees were cut and held to determine emergence rate of A. glabripennis. Unique marks were painted onto 912 beetles released into a group of 165 trees in Gansu, China. Data on subsequent sightings of beetles were used in a truncated diffusion model to calculate flight distances. Characteristics of the trees and climatic information were used in statistical tests for influence on movement. A total of 2245 sightings of beetles were observed and 29% of marked beetles were resighted. The scanning technique using binoculars was 90% effective in finding beetles and provided 81% accuracy for determining the sex of the beetles. Experimental manipulation of density quantified how A. glabripennis congregated on unoccupied trees and were repulsed from crowded hosts. The seasonal emergence rate of adults declined exponentially from July 20 to August 5. The results suggested A. glabripennis fly to nearby host trees at a rate of 34% per day. Median flight distance was estimated at 20 m per day. Statistical analysis with a generalized linear model tested the beetle's propensity to leave a tree and distance of flight. Generally, beetle movement showed a significant response to beetle density, weather conditions, beetle size, and tree size, in that order. The techniques developed here improve on previous recapture techniques to quantify dispersal and can be useful for analysing populations of other organisms.     

Physiological development and dispersal ability of newly emerged Monochamus galloprovincialis

Novel associations between exotic pathogens and native insects may result in major ecological and economical losses. In Europe, Monochamus galloprovincialis (Olivier) (Coleoptera: Cerambycidae) is the only known vector of the exotic pine wood nematode Bursaphelenchus xylophilus (Steiner & Buhrer) Nickle (Nematoda: Aphelenchoididae), the causal agent of pine wilt disease (PWD). Transportation of goods containing nematode‐infested beetles is the main pathway for the spread of the disease. In this scenario, management actions involving early detection and eradication are critical to stop further spread by the vector. Although dispersal of mature M. galloprovincialis has been successfully tracked using commercial baits and traps, dispersal ability of immature individuals is poorly understood. Sexual maturation and other physiological traits related to dispersal were studied for newly emerged M. galloprovincialis after different shoot‐feeding spans. Sexual maturation was complete after on average 12 (males) or 13 (females) feeding days. Monochamus galloprovincialis adults emerged with an average of 10% lipid and 23.8% (males) or 29.9% (females) pterothorax muscle content, and these percentages did not change significantly during shoot feeding. Microtomography images of wing muscle structures at different maturation stages confirmed these results. Emerged adults that were kept unfed survived an average of 12 days in the lab. The ability of unfed immature insects to fly through hostless terrain was studied by marking and releasing newly emerged insects from a crop area located up to 3 km away from two small pine stands. The longest flown distance recorded was 2 km. Fitted regressions indicated that immature insects could have travelled up to 3 109 m to reach the experimental stands. We found that M. galloprovincialis emerge with well‐developed thoracic muscles and energy reserves that enable them to sustain long flights over non‐forested areas. These findings should aid managers and policy makers in devising sound procedures in areas where the risk of introducing PWD is high.     

Limited dispersal and its effect on population structure in the milkweed beetle Tetraopes tetraophthalmus

Summary The movement patterns of adult milkweed beetles, Tetraopes tetraphthalmus, were monitored via a mark-recapture technique. Movement or dispersal patterns were studied in two natural populations, one in which the host plant, Asclepias syriaca, was nearly continuously distributed over a 250×90 m area and another where Asclepias was distributed in 17 small discrete patches. In both populations dispersal distances resulting from the flight patterns of the adult beetles were quite short, averaging less than 40 m from the point of first encounter 10 days after marking. Males were shown to be more vagile than females. The distribution of dispersal distances collected from one of the populations was fit to three statistical distributions cited in the literature as expected from dispersal by many small-scale movements or observed in other species. It was found that an equation describing an exponential decay gave the best statistical fit to the data collected here for milkweed beetles. The data is discussed in the context of the effects of the limited dispersal power of the beetles and the distribution of suitable habitat on the population structure of Tetraopes.     

Lethal and sublethal effects of thiacloprid on non-target carpenter ant,Camponotus japonicus Mayr (Hymenoptera: Formicidae)

To control population of Monochamus beetles that transmit pine wood nematode, Bursaphelenchus xylophilus, a variety of insecticides have been applied to forest ecosystems in Korea. Non-target predatory insects can be directly or indirectly exposed to insecticides. We evaluated potential lethal and sublethal effects of thiacloprid on survival and behavior of carpenter ants, Camponotus japonicus Mayr. Field-collected ants were directly exposed to several food items such as thiacloprid-exposed Monochamus beetles, 10% sugar water with thiacloprid concentrations, and 10% sugar water at group and individual levels. In experiment for groups of individuals, dead beetle bodies generally had possible adverse effects on ants through dietary exposure, because two forager ants were dead or paralysis after they were exposed to thiacloprid-exposed Monochamus beetles. At individual level, dietary exposure to thiacloprid at concentrations of 10 and 50?mg/L was lethal to ants, causing paralysis and impaired walking. Mortality of ant workers was higher in direct or indirect exposure treatments than that in the control. Some ants exposed to thiacloprid showed abnormal behavior within a few days, especially at thiacloprid concentrations of 10 and 50?mg/L. However, some of them sometimes recovered from the abnormal behavior in a day. In consequence, application of thiacloprid in pine forests may disrupt species interaction and foraging behavior of ants, but the effect of thiacloprid through trophallaxis behavior should be further assessed using more extensive colonies composed of queen, workers, brood, and so on.     

Inter-population dispersal by adult stoneflies detected by stable isotope enrichment

1. Dispersal of adult stream insects may be of considerable importance in regional population dynamics and colonisation of new sites, but quantifying the rate and extent of dispersal is difficult. 2. We used stable isotope (¹⁵N) enrichment to mark more than 1.5 million larval stoneflies (Leuctra inermis) before they emerged from an upland stream in the Plynlimon area of mid‐Wales, in order to determine directly the rate and pattern of inter‐site dispersal. 3. A small number of isotopically enriched adult stoneflies were captured in samples taken at adjacent streams between 800 m and 1.1 km away from the source population, including a headwater of a different river system. 4. The distribution of marked individuals suggested that wind influences dispersal direction in the uplands, but the low number of captures limits our ability to draw firm conclusions. 5. This is the first direct demonstration of dispersal of insects between streams. The dispersal distances recorded were significantly greater than those suggested by previous direct studies, but much more consistent with indirect studies based on genetic differentiation of populations.     

The role of herbivory by wood-boring insects in mangrove ecosystems in Belize

The roles wood-boring insects play in modifying mangrove ecosystems were examined on small, offshore mangrove islands in Belize. Several species of xylem- and phloem-feeding woodborers consume the wood of living mangrove trees. By girdling, pruning, and hollowing, woodborers killed over 50% of the Rhizophora mangle canopy in experimental plots arrayed across a tidal-elevation gradient. In contrast, leaf-feeding herbivores removed less than 6% of the canopy. In the plots, stem girdlers killed over three branches per tree. The patterns of herbivory by three functional feeding groups were heterogeneous and did not vary consistently with tidal elevation. Because R. mangle lacks viable axillary buds or the ability to produce epicormic shoots to replace pruned branches, the canopy architecture was significantly modified by this damage. The branches that were pruned by stem girdlers created numerous small holes or gaps in the mangrove canopy. Shoot growth and flowering increased in R. mangle trees with 50% of their branches experimentally girdled. Because branches and twigs attacked by woodborers lost their leaves prematurely as greenfall, the quantity and quality of leaf litter were altered when a leaf-bearing branch was girdled or hollowed. These changes suggest that wood-boring insects also significantly affect internal and external nutrient cycling processes in mangrove ecosystems.     

Bark beetle and woodborer responses to stand thinning and prescribed fire in northeastern US coastal and inland pitch pine barrens

1. Southern pine beetle, Dendroctonus frontalis, has expanded its range further into the northeastern United States. This expansion threatens rare and ecologically valuable interior and coastal pitch pine barrens.
2. Pitch pine barrens restoration and southern pine beetle infestation suppression often involve leaving downed dead wood that saproxylic insects can exploit.
3. Semiochemical-baited traps were used to investigate the response of bark beetles and woodborers to restoration treatments at Rocky Point State Forest and the Albany Pine Bush Preserve, examples of coastal and interior pitch pine forests, respectively.
4. A total of 29,598 saproxylic insects from 116 species of bark beetles and woodborers were captured at Rocky Point State Forest, while 23,117 individuals from 67 species were captured at Albany Pine Bush Preserve.
5. Ips spp. were abundant at both sites with 28%–47% and 42%–74% of total collections at Rocky Point State Forest and Albany Pine Bush Preserve, respectively.
6. Ips grandicollis did not respond to treatments at either site. However, Ips pini was found in higher numbers in thinned blocks in Rocky Point State Forest.

     

Isolation and characterisation of Isaria farinosa and Purpureocillium lilacinum associated with emerald ash borer,Agrilus planipennis in Canada

Entomopathogenic fungi of the genera Isaria and Purpureocillium were recovered from infestation sites of emerald ash borer (EAB) in Southern Ontario, Canada. Isolates were identified using morphological characters and by sequencing the ITS1-5.8S-ITS2 ribosomal DNA gene and partial β-tubulin gene. Phylogenetic analysis and constructed trees based on the ITS and β-tubulin gene explicitly confirm isolates L66B, SY17-a and LHY46-a as Isaria farinosa and B3A, B59A and SY45B-a as Purpureocillium lilacinum. Pathogenicity was tested in the laboratory against adult EAB using a single concentration (2.0×10⁷ conidia/ml) applied topically to adults. Controls included three commercial isolates: Isaria fumosorosea LRC176, Metarhizium brunneum LRC187 and Beauveria bassiana strain GHA. The native isolates I. farinosa L66B and P. lilacinum SY45B-a killed 75 and 50% of the beetles 14 days post-inoculation. Although these indigenous entomogenous fungi were less virulent compared with the commercial isolates, yearly isolation from EAB populations suggests they are one of the natural mortality factors of EAB in Canada.     

Emergence patterns of univoltine and bivoltine Ips typographus (L.) populations and associated natural enemies

Control measures aiming at reducing bark beetle populations and preserving their natural enemies require a sound knowledge on their overwintering and emergence behaviour. These behavioural traits were investigated in univoltine and bivoltine populations of the European spruce bark beetle (Ips typographus [L.], Coleoptera: Scolytinae) and its predators and parasitoids over several consecutive years. In univoltine populations, roughly 50% of the bark beetles left their brood trees in fall together with most parasitoids and some significant predatory flies and beetles. In bivoltine populations, <10% of the second bark beetle generation emerged in fall and the remainder overwintered under the bark of their brood trees. Likewise, most predatory beetles and flies spent wintertime with their prey under the bark, while most parasitic wasps emerged in fall. The spring emergence of bivoltine predatory beetles was found to occur up to 3 weeks earlier than that of I. typographus, while that of the predatory flies and the parasitoids was delayed by up to 1 month. In univoltine populations, the bark beetles emerged several weeks prior to most antagonistic taxa. In the heat year 2003, three I. typographus generations were produced at the lower location, 36% of the third generation emerged in fall, while the proportions of overwintering predators remained largely the same as in previous years. Similar to their host, more parasitoids left their brood trees in fall after warm years. The results show that sanitation felling during winter probably kills most bark beetles in bivoltine populations, but also eliminates many natural enemies. In univoltine populations, sanitation felling might be less detrimental to both I. typographus and natural enemies because a fair fraction of their populations will already have left the trees before cutting. Warmer climates may affect the interactions of bark beetles and natural enemies and thus the impact of control measures.